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PubMed2006

Cell behaviors associated with somite segmentation and rotation in Xenopus laevis.

During vertebrate development the formation of somites is a critical step, as these structures will give rise to the vertebrae, muscle, and dermis. In Xenopus laevis, somitogenesis consists of the partitioning of the presomitic mesoderm into somites, which undergo a 90-degree rotation to become aligned parallel to the notochord. Using a membrane-targeted green fluorescent protein to visualize cell outlines, we examined the individual cell shape changes occurring during somitogenesis. We show that this process is the result of specific, coordinated cell behaviors beginning with the mediolateral elongation of cells in the anterior presomitic mesoderm and then the subsequent bending of these elongated cells to become oriented parallel with the notochord. By labeling a clonal population of paraxial mesoderm cells, we show that cells bend around their dorsoventral axis. Moreover, this cell bending correlates with an increase in the number of filopodial protrusions, which appear to be posteriorly directed toward the newly formed segmental boundary. By examining the formation of somites at various positions along the anteroposterior axis, we show that the general sequence of cell behaviors is the same; however, somite rotation in anterior somites is slower than in posterior somites. Lastly, this coordinated set of cell behaviors occurs in a dorsal-to-ventral progression within each somite such that cells in the dorsal aspect of the somite become aligned along the anteroposterior axis before cells in other regions of the same somite. Together, our data further define how these cell behaviors are temporally and spatially coordinated during somite segmentation and rotation.

Animals↗
PubMed1997

Regulation of paraxis expression and somite formation by ectoderm- and neural tube-derived signals.

During vertebrate embryogenesis, the paraxial mesoderm becomes segmented into somites, which form as paired epithelial spheres with a periodicity that reflects the segmental organization of the embryo. As a somite matures, the ventral region gives rise to a mesenchymal cell population, the sclerotome, that forms the axial skeleton. The dorsal region of the somite remains epithelial and is called dermomyotome. The dermomyotome gives rise to the trunk and limb muscle and to the dermis of the back. Epaxial and hypaxial muscle precursors can be attributed to distinct somitic compartments which are laid down prior to overt somite differentiation. Inductive signals from the neural tube, notochord, and overlying ectoderm have been shown to be required for patterning of the somites into these different compartments. Paraxis is a basic helix-loop-helix transcription factor expressed in the unsegmented paraxial mesoderm and throughout epithelial somites before becoming restricted to epithelial cells of the dermomyotome. To determine whether paraxis might be a target for inductive signals that influence somite patterning, we examined the influence of axial structures and surface ectoderm on paraxis expression by performing microsurgical operations on chick embryos. These studies revealed two distinct phases of paraxis expression, an early phase in the paraxial mesoderm that is dependent on signals from the ectoderm and independent of the neural tube, and a later phase that is supported by redundant signals from the ectoderm and neural tube. Under experimental conditions in which paraxis failed to be expressed, cells from the paraxial mesoderm failed to epithelialize and somites were not formed. We also performed an RT-PCR analysis of combined tissue explants in vitro and confirmed that surface ectoderm is sufficient to induce paraxis expression in segmental plate mesoderm. These results demonstrate that somite formation requires signals from adjacent cell types and that the paraxis gene is a target for the signal transduction pathways that regulate somitogenesis.

Amino Acid Sequence↗
PubMed2004

Mutations affecting somite formation in the Medaka (Oryzias latipes).

The metameric structure of the vertebrate trunk is generated by repeated formation of somites from the unsegmented presomitic mesoderm (PSM). We report the initial characterization of nine different mutants affecting segmentation that were isolated in a large-scale mutagenesis screen in Medaka (Oryzias latipes). Four mutants were identified that show a complete or partial absence of somites or somite boundaries. In addition, five mutations were found that cause fused somites or somites with irregular sizes and shapes. In situ hybridization analysis using specific markers involved in the segmentation clock and antero-posterior (A-P) polarity of somites revealed that the nine mutants can be compiled into two groups. In group 1, mutants exhibit defects in tailbud formation and PSM prepatterning, whereas A-P identity in the somites is defective in group 2 mutants. Three mutants (planlos, pll; schnelles ende, sne; samidare, sam) have characteristic phenotypes that are similar to those in zebrafish mutants affected in the Delta/Notch signaling pathway. The majority of mutants, however, exhibit somitic phenotypes distinct from those found in zebrafish, such as individually fused somites and irregular somite sizes. Thus, these Medaka mutants can be expected to provide clues to uncovering novel components essential for somitogenesis.

Animals↗
PubMed1996

Spatial and temporal effects of axial structures on myogenesis of developing somites.

To elucidate the precise roles of axial structures in the myogenic differentiation of the somite, we have examined the effects of the axial organs' precise spatial position during migration and differentiation of somitic cells by using in vivo transplantation of the neural tube and of the notochord directly into the paraxial mesoderm. Differentiation of myotomal cells was identified through the use of Quox 1 antibody which recognizes specifically a quail homeoprotein Quox 1. We have demonstrated that both ectopic neural tube and notochord are able to influence the myogenesis in somites, but that the spatial position of axial organs and the degree of somite maturation at grafting time are decisive. At the level of the somites which were already formed and developmentally advanced (somites III-VI), both neural tube and notochord promote myogenesis, and the promoting effect of notochord is more efficient than that of the neural tube. In the newly formed somites (I-II) and/or the segmental plate mesoderm, the notochord inhibits the myogenesis of somites, whereas the neural tube plays an evident myogenic promoting role. But the myogenic effect of the neural tube depends not only upon the stage of developing somites and presomitic mesoderm, but also on the developmental maturation of the neural tube. We have demonstrated that the myogenic effect of the rostral part of neural tube is stronger than that of its caudal part. This observation suggests that there is a gradient of myogenic effect along the rostrocaudal axis of the neural tube, which depends on the developmental maturation of neural tube, and that the generation of skeletal muscle during somitogenesis may be in relation with the rostrocaudal gradient of the capacity of the neural tube to stimulate myogenesis since somites are also distributed along an anteroposterior axis.

Animals↗
PubMed1998

Mediolateral patterning of somites: multiple axial signals, including Sonic hedgehog, regulate Nkx-3.1 expression.

The axial structures, the notochord and the neural tube, play an essential role in the dorsoventral patterning of somites and in the differentiation of their many cell lineages. Here, we investigated the role of the axial structures in the mediolateral patterning of the somite by using a newly identified murine homeobox gene, Nkx-3.1, as a medial somitic marker in explant in vitro assays. Nkx-3.1 is dynamically expressed during somitogenesis only in the youngest, most newly-formed somites at the caudal end of the embryo. We found that the expression of Nkx-3.1 in pre-somitic tissue explants is induced by the notochord and maintained in newly-differentiated somites by the notochord and both ventral and dorsal parts of the neural tube. We showed that Sonic hedgehog (Shh) is one of the signaling molecules that can reproduce the effect of the axial structures by exposing explants to either COS cells transfected with a Shh expression construct or to recombinant SHH. Shh could induce and maintain Nkx-3.1 expression in pre-somitic mesoderm and young somites but not in more mature, differentiated ones. The effects of Shh on Nkr-3.1 expression were antagonized by a forskolin-induced increase in the activity of cyclic AMP-dependent protein kinase A. Additionally, we confirmed that the expression of the earliest expressed murine myogenic marker, myf 5, is also regulated by the axial structures but that Shh by itself is not capable of inducing or maintaining it. We suggest that the establishment of somitic medial and lateral compartments and the early events in myogenesis are governed by a combination of positive and inhibitory signals derived from the neighboring structures, as has previously been proposed for the dorsoventral patterning of somites.

Animals↗
PubMed2004

The T-box transcription factor Tbx18 maintains the separation of anterior and posterior somite compartments.

The compartmentalization of somites along their anterior-posterior (AP) axis is pivotal to the segmental organization of the vertebrate axial skeleton and the peripheral nervous system. Anterior and posterior somite halves contribute to different vertebral elements. They are also characterized by different proliferation rates and properties with respect to neural crest cell migration and spinal nerve passage. AP-somite polarity is generated in the anterior presomitic mesoderm by Mesp2 and Delta/Notch signaling. Here, we demonstrate that maintenance of AP-somite polarity is mediated by the T-box transcription factor Tbx18. Mice deficient for Tbx18 show expansion of pedicles with transverse processes and proximal ribs, elements derived from the posterior lateral sclerotome. AP-somite polarity is established in Tbx18 mutant embryos but is not maintained. During somite maturation, posterior somite compartments expand most likely because of posterior cells invading the anterior somite half. In the anterior lateral sclerotome, Tbx18 acts as an antiapoptotic factor. Ectopic expression experiments suggest that Tbx18 can promote anterior at the expense of posterior somite compartments. In summary, Tbx18 appears to act downstream of Mesp2 and Delta/Notch signaling to maintain the separation of anterior and posterior somite compartments.

Animals↗
PubMed2002

Signals that instruct somite and myotome formation persist in Xenopus laevis early tailbud stage embryos.

Somite formation is a lengthy process that begins at gastrulation and continues through tailbud stages to form approximately 50 pairs of somites in the frog, Xenopus laevis. In Xenopus, the somite primarily gives rise to myotome. We sought to determine whether the formation of somites and myotome requires a transient signal active during gastrulation or a constitutive signal active throughout development to instruct dorsal mesodermal cells to form the posterior somites. Previous work from our lab revealed that cells from the neural ectoderm are capable of responding to mesoderm-inducing signals [Domingo and Keller: Dev Biol 2000;225:226-240]. Thus, to test for the presence of somite-inducing signals, we performed a series of grafting experiments in which we used gastrula cells from the anterior neural ectoderm (ANE). Fluorescently labeled ANE cells were grafted to the posterior paraxial mesoderm of progressively older host embryos between stages 11 (mid gastrula) and 23 (early tailbud). Our results showed that signals within the paraxial mesoderm can instruct prospective ANE cells, which normally give rise to head structures, to instead differentiate into myotome cells. We found that the grafted cells adopted the local paraxial mesoderm cell behaviors, which consists of mediolateral intercalation, segmentation, somite cell rotation, and differentiation to myotome. In addition, we show that the grafted ANE cells that adopt a myotome morphology also express the muscle-specific marker, 12/101. Through a series of heterochronic grafts, we determined that the duration of somite-inducing signals extends from the early gastrula (stage 11) through the early tailbud (stage 23) stage embryos. These results demonstrate that somite induction is not a transient event that occurs during gastrulation, but that it is instead a continuous event that can occur as new somites are added to the posterior axis.

Animals↗
PubMed2006

Gli3-mediated somitic Fgf10 expression gradients are required for the induction and patterning of mammary epithelium along the embryonic axes.

Little is known about the regulation of cell fate decisions that lead to the formation of five pairs of mammary placodes in the surface ectoderm of the mouse embryo. We have previously shown that fibroblast growth factor 10 (FGF10) is required for the formation of mammary placodes 1, 2, 3 and 5. Here, we have found that Fgf10 is expressed only in the somites underlying placodes 2 and 3, in gradients across and within these somites. To test whether somitic FGF10 is required for the formation of these two placodes, we analyzed a number of mutants with different perturbations of somitic Fgf10 gradients for the presence of WNT signals and ectodermal multilayering, markers for mammary line and placode formation. The mammary line is displaced dorsally, and formation of placode 3 is impaired in Pax3ILZ/ILZ mutants, which do not form ventral somitic buds. Mammary line formation is impaired and placode 3 is absent in Gli3Xt-J/Xt-J and hypomorphic Fgf10 mutants, in which the somitic Fgf10 gradient is shortened dorsally and less overall Fgf10 is expressed, respectively. Recombinant FGF10 rescued mammogenesis in Fgf10(-/-) and Gli3Xt-J/Xt-J flanks. We correlate increasing levels of somitic FGF10 with progressive maturation of the surface ectoderm, and show that full expression of somitic Fgf10, co-regulated by GLI3, is required for the anteroposterior pattern in which the flank ectoderm acquires a mammary epithelial identity. We propose that the intra-somitic Fgf10 gradient, together with ventral elongation of the somites, determines the correct dorsoventral position of mammary epithelium along the flank.

Animals↗
PubMed1995

Migration of myogenic cells from the somites to the fore-limb buds of developing mouse embryos.

In this study, we have isolated newly formed somites from the caudal regions of 8.5 day mouse embryos and transplanted them orthotopically into correspondingly staged hosts at the level of the prospective limb-forming region. The experimental embryos were then cultured intact for 32-36 hr. The donor somites used were pre-labelled with DiI, a fluorescent lipophilic dye, or were obtained from transgenic embryos that carried a 1 kb 5' regulatory sequence of the desmin gene linked to the gene encoding Escherichia coli beta-galactosidase. The transgene is specifically expressed in skeletal muscles (Li et al. [1993] Development 117:947-959). The aim of these experiments was to show definitively that the musculature of the mammalian limb is derived from the somites. The results demonstrated that DiI-labelled cells from the implanted somites were able to invade the proximal region of the fore-limb bud during the course of development. The use of transgenic somites as grafts confirmed that some of the somitic cells found in the limbs were myogenic cells. To determine whether the displacement of somitic cells is an active or passive process, somatopleure obtained from the prospective limb-forming regions of day 8.5 day embryos was implanted into 8.5 day hosts. We did not detect the presence of DiI-labelled somatopleural cells in the fore-limb after 32-36 hr of culture. This suggests that somitic cells reached the limb bud via active locomotion rather than as a result of being passively dragged there, as the limb elongates during development. In addition, we injected latex beads into the somites, as probes, to determine whether extracellular matrix-driven translocation plays a role in driving the somitic cells to the limb bud. In a majority of the specimens examined, we could not detect the presence of these beads in the limb bud. However, in the trunk of these embryos, the beads were found dispersed throughout the ventral neural crest pathway.

Animals↗
PubMed1991

The effect of heat shocks, which alter somite segmentation, on Rohon-Beard neurite outgrowth from the spinal cord of Xenopus embryos.

The vertebrate spinal cord shows a segmental pattern of dorsal and ventral nerve roots. It is believed that this peripheral neural segmentation depends on the segmentation of the somitic mesoderm into somites. I have studied the relationship between somite segmentation and the pattern of Rohon-Beard sensory neurite outgrowth from the spinal cord in Xenopus embryos. Most Rohon-Beard neurites grow out in fascicles from the spinal cord at the intersomite clefts, a smaller number of neurites growing out over the dorsal somite margins. Previous work on peripheral neural segmentation has relied on transplanting or destroying somitic mesoderm. Here a non-invasive technique has been used. The pattern of somite segmentation was disrupted by heat shocks, leading to regions of somite fusion where one or two intersomite clefts are absent. At the regions of somite fusion the number of sensory fascicles is then unchanged but their distribution is different. The segmental pattern of sensory fascicle outgrowth is replaced by a fairly even distribution of outgrowth fascicles. Two interpretations of the difference in fascicle outgrowth at fused and control somites are discussed. Firstly, that heat shocks have a direct affect on fascicle outgrowth. Secondly, that heat shocks affect fascicle outgrowth indirectly by disrupting somite segmentation.

Animals↗
PubMed1988

The effect of somite manipulation on the development of motoneuron projection patterns in the embryonic chick hindlimb.

Although the formation of motoneuron projections to individual muscles in the embryonic chick hindlimb has been shown to involve the specific recognition of environmental cues, the source of these cues and their mode of acquisition are not known. I show in the accompanying paper (C. Lance-Jones, 1988, Dev. Biol. 126, 394-407) that there is a correlation between the segmental level of origin of motoneurons and the somitic level of origin of the muscle cells of their targets in the chick hindlimb. These data are compatible with the hypothesis that the developmental basis for specific recognition is a positional one. Motoneurons and myogenic cells may be uniquely labeled in accord with their axial level of origin early in development and subsequently matched on the basis of these labels. To test this hypothesis, I have assessed motoneuron projection patterns in the embryonic chick hindlimb after somitic tissue manipulations. In one series of embryos, somitic mesoderm at levels 26-29 or 27-29 was reversed about the anteroposterior axis prior to myogenic cell migration and axon outgrowth. Since previous studies have shown that cells migrate from the somites in accord with their position and that somites 26-29 populate anterior thigh musculature, this operation will have reversed the somitic level of origin of anterior thigh muscles. Retrograde HRP labeling of projections to anterior thigh muscles at stage (st) 30 and st 35-38 showed that motoneuron projections were largely normal. This finding suggests that limb muscle cells or their source, the somites, do not contain the cues responsible for specific recognition prior to myogenic cell migration and axon outgrowth. To confirm that specific guidance cues were still intact after somitic mesoderm reversal, I also assessed motoneuron projections in embryos where somitic tissue plus adjacent spinal cord segments at levels 26-29 were reversed in a similar manner. Analyses of the distribution of retrogradely labeled motoneurons in reversed cord segments at st 35-36 indicated that motoneuron projections were reversed. This finding suggests that motoneurons have altered their course to project to correct targets despite the altered somitic origin of their targets and, thus, that specific guidance cues were intact. I conclude that if cues governing target or pathway choice are encoded positionally then they must be associated with other embryonic tissues such as the connective tissues or that guidance cues are acquired by myogenic cells after the onset of migration and motoneuron specification.

Animals↗
PubMed1996

Chicken Pax-1 gene: structure and expression during embryonic somite development.

Recent mouse genetic studies have implicated Pax-1, a paired-box-containing gene, in sclerotomal differentiation and vertebral body formation. To investigate Pax-1 function in somitic sclerotomal differentiation in the chick embryo, we have cloned the chicken Pax-1 gene, and its full length cDNA, and characterized its temporal and spatial expression pattern during somite development. Sequence analysis shows that chicken Pax-1 is highly homologous to murine and human Pax-1 genes with respect to the putative DNA-binding paired-box domain and the octapeptide domain. Northern analysis using probes derived from the paired-box domain and a unique non-paired box sequence of chicken Pax-1 detected 2-kb mRNA transcript. The expression profiles of Pax-1 were examined by in situ hybridization and Northern analysis. The first detectable expression of Pax-1 is seen in the most caudal epithelial somite. As the somite matures, Pax-1 expression takes on a medial distribution, thus corresponding to but preceding the emergence of the sclerotome. In the more mature, rostral somites (stage V and older), Pax-1 expression is found to be progressively localized first to the ventral-medial regions, and then to the caudal-ventral-medial quadrant of the mature somite. This pattern strongly supports the notion that Pax-1 expression is involved in somitogenesis and sclerotomal differentiation, and that it is subsequently a characteristic of the caudal half of the sclerotome, the presumptive precursor of vertebral cartilage. Northern analysis substantiated this expression profile and further revealed that the level of somitic Pax-1 expression increases as a function of embryonic development. Finally, we subjected chicken embryos to controlled heat shock treatment to perturb somite formation and segmentation. The pattern of Pax-1 expression in the anomalous somitic structures generated by controlled heat shock further supports a functional role for Pax-1 in somite development.

Amino Acid Sequence↗
PubMed1996

A quantitative study of the segmental distribution of somitic cells in the developing chick limb bud using laser-scanning confocal microscopy.

Our aim was to map the segmental distribution of somitic cells in the limb before and after the fusion of these cells into myotubes. Somitic cells of the brachial somites were labeled by injection of DiI and DiO into the somitoceles of embryonic Day 2 (E2) embryos. The quantitative distribution of dye-labeled cells from injected somites was examined in whole mounts of E4 wing buds and in sections of E5 wing buds using confocal microscopy. Dye derived from cells of anterior brachial somites 16 and 17 was highly concentrated in the anterior half of E4 wings, whereas dye from posterior brachial somites 20 and 21 was concentrated in the posterior half of the wing. Not more than 14% (on average) of the dye in the wing was found outside the preferred half. Dye was equally dispersed in the anterior and posterior halves of the wing when the middle two somites contributing to the wing musculature, numbers 18 and 19, were injected. The total amount of dye in the whole limb at E4 was not significantly different after injection of similar amounts of dye into either pair of brachial somites 16/17, 18/19, or 20/21. In E5 embryos the distribution of labeled cells in the newly formed muscle masses, identified with antibody 2H2 to myosin light chains, was examined in cryostat cross-sections of the wing. Dye was more widely distributed in the wing than at E4. As much as 32% (on average) of the dye was found in the muscle mass outside the preferred half after administration of dye into the anterior or posterior pair of somites. We conclude that the branchial somites contribute similar numbers of cells to the wing musculature and that the segmental origin of these cells is rigidly maintained during their lateral migration into the limb. Cells from adjacent segments then "mix" when they fuse to form myotubes at E5. In addition, we found a significantly greater amount of dye per unit of muscle mass in the proximal compared with distal parts of the limb. We argue that this finding is due to division of myogenic cells during the course of their migration in the wing.

Animals↗
PubMed1988

The allocation of cells in the presomitic mesoderm during somite segmentation in the mouse embryo.

Orthotopic grafts of wheat germ agglutinin-colloidal gold conjugate (WGA-gold) labelled cells were used to demonstrate differences in the segmental fate of cells in the presomitic mesoderm of the early-somite-stage mouse embryos developing in vitro. Labelled cells in the anterior region of the presomitic mesoderm colonized the first three somites formed after grafting, while those grafted to the middle region of this tissue were found mostly in the 4th-7th newly formed somites. Labelled cells grafted to the posterior region were incorporated into somites whose somitomeres were not yet present in the presomitic mesoderm at the time of grafting. There was therefore an apparent posterior displacement of the grafted cells in the presomitic mesoderm. Colonization of somites by WGA-gold labelled cells was usually limited to two to three consecutive somites in the chimaera. The distribution of cells derived from a single graft to two somites was most likely due to the segregation of the labelled population when cells were allocated to adjacent meristic units during somite formation. Further spreading of the labelled cells to several somites in some cases was probably the result of a more extensive mixing of mesodermal cells among the somitomeres prior to somite segmentation.

Animals↗
PubMed1993

Delayed somite formation in a quail line exhibiting myofiber hyperplasia is accompanied by delayed expression of myogenic regulatory factors and myosin heavy chain.

A myofiber hyperplastic quail line P has been developed through selection for heavy body weight. Since the number of muscle fibers is determined early in development and skeletal muscle originates from somites, we compared somite formation and muscle-specific gene expression in P- and control C-line quail embryos. At 47 hours of incubation, C embryos had 18 somite pairs and P embryos had 14.3. By 72 and 120 hours, both lines appeared to be at the same stage of somite development. To determine whether the delay in the formation of the brachial somites was accompanied by alterations in muscle-specific gene expression, we conducted whole-mount in situ hybridization and immunofluorescence studies. At 47 hours of incubation, C embryos were expressing qmf1 in the first 12 somites, while in P embryos only the first 7 somites showed qmf1 activation. Delays in expression were also observed for qmf3 at 43 hours and for all three myogenic factors (qmf1, qmf2 and qmf3) at 60 hours. At 65 hours, C embryos expressed myosin heavy chain in the first 15 somite pairs and P embryos in the first 7. At 72 hours, the transient delay in somite formation had disappeared and there was no lag in myosin heavy chain expression between the lines. The phase delay in brachial somite formation, myogenic factors and myosin heavy chain expression may be associated with the observed myofiber hyperplasia in P-line quail by allowing an increase in the muscle stem cell population.

Animals↗
PubMed1979

The mechanism of somite segmentation in the chick embryo.

The segmentation of somites in the chick embryo has been studied by transmission and scanning electron microscopy (stages 8-14). The segmental plate mesoderm consists of loosely arranged mesenchymal cells, whereas the newly formed somites are composed of elongated, spindle-shaped cells arranged radially around a lumen, the myocoele. The diamter of each somite is thus two cells plus the myocoele. Two major factors appear to be responsible for the change in cell shape at segmentation: (1) Each prospective somite cell becomes anchored at one end to the adjacent epithelia (i.e. the neural tube, the notochord, the ectoderm, the endoderm or the aorta) by means of collagen fibrils. These fibrils are already present in the segmental plate before the somites begin to form. (2) A change in cell-to-cell adhesiveness causes the free ends of these cells to adhere to one another. (Bellairs, Curtis & Sanders, 1978). This adhesion is then supplemented by the development of tight junctions proximally in the somite. Because it is anchored at both ends, each somite cell is under tension in much the same way as a fibroblast cell in tissue culture is under tension. Each somite cell therefore becomes elongated and the somite as a whole accommodates its general shape to that of the space available between the adjacent tissues. The arrangement of the cells in the more differentiated somites (stages 17-18) has also been examined and it has been found that the chick resembles Xenopus in that the myotome cells undergo rotation and become orientated in an anteroposterior direction.

Animals↗
PubMed2006

Asymmetric localization of numb in the chick somite and the influence of myogenic signals.

Whereas Notch signaling is known to play an essential role in the formation of somites, its role during later stages of somite maturation is less well understood. Here, we examine the signals and transcription factors that control the expression of the Notch antagonist, Numb, during somite maturation in the chick embryo. Numb mRNA is present in the epithelial somite and is increased in expression in the forming myotome. Numb protein displays a very specific subcellular localization and dynamic expression during somite maturation. Numb protein is asymmetrically localized in a cortical crescent on the basal side of dividing cells in the dorsomedial lip of the dermomyotome and is subsequently uniformly distributed throughout differentiated myotomal cells. Treatment of somites with either the combination of Wnt-3a and Shh, or ectodermal signals plus noggin, both of which induce somitic myogenesis, did not significantly affect Numb transcript levels but did lead to a dramatic increase in the levels of Numb protein, which was uniformly distributed throughout the cytoplasm of the resultant myotubes. Forced expression of MyoD in somites similarly induced high levels of Numb protein throughout the cytoplasm, without affecting Numb mRNA levels. We also found that signals that promote somitic myogenesis or forced MyoD expression induced expression of the Notch ligand, Serrate-2. Our findings suggest that Notch signals are specifically repressed in the myotome and that asymmetric expression of Numb in dividing cells of the dorsomedial lip of the dermomyotome may modulate whether these cells continue to divide or differentiate into myotomal cells.

Animals↗
PubMed1997

The fate of the first avian somite.

We have studied the derivatives of the first somite using the quail-chick marking technique. After transplantation of the somite, the chick embryos were reincubated for periods ranging from 4 h to 11 days. Coronal and sagittal sections of the embryos were prepared for parallel staining with Feulgen-reaction, anti-quail antibody, anti-desmin antibody and QH-1 antibody. The first somite consists of an epithelial envelope surrounding somitocoele cells. Like other somites, it forms sclerotome, dermatome and myotome. Cells contribute to the occipital and parasphenoid bone, the meninges, the dermis in the occipital region and the pharyngeal connective tissue. The contribution of the first somite to bones, meninges, dermis and pharyngeal connective tissue is characterised by sharp anterior and posterior boundaries. In contrast, other derivatives such as connective tissue surrounding the vagus nerve, the carotid artery, and jugular vein exceed 10 to 18 segments. This is also true for myogenic cells participating in the formation of the cucullaris capitis muscle that extends from the temporal bone to the shoulder. In one third of the embryos, myocytes of the intrinsic laryngeal muscles are derived from the grafted first somite. Moreover, endothelial cells originate from this somite and migrate into the head (hind-brain, meninges, dermis), neck (pharynx, connective tissue surrounding the vagus nerve, carotid artery and jugular vein) and thorax. With respect to differentiation and derivatives the first somite is similar to other somites.

Animals↗