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PubMed2004

Eya1 and Six1 are essential for early steps of sensory neurogenesis in mammalian cranial placodes.

Eya1 encodes a transcriptional co-activator and is expressed in cranial sensory placodes. It interacts with and functions upstream of the homeobox gene Six1 during otic placodal development. Here, we have examined their role in cranial sensory neurogenesis. Our data show that the initial cell fate determination for the vestibuloacoustic neurons and their delamination appeared to be unaffected in the absence of Eya1 or Six1 as judged by the expression of the basic helix-loop-helix genes, Neurog1 that specifies the neuroblast cell lineage, and Neurod that controls neuronal differentiation and survival. However, both genes are necessary for normal maintenance of neurogenesis. During the development of epibranchial placode-derived distal cranial sensory ganglia, while the phenotype appears less severe in Six1 than in Eya1 mutants, an early arrest of neurogenesis was observed in the mutants. The mutant epibranchial progenitor cells fail to express Neurog2 that is required for the determination of neuronal precursors, and other basic helix-loop-helix as well as the paired homeobox Phox2 genes that are essential for neural differentiation and maintenance. Failure to activate their normal differentiation program resulted in abnormal apoptosis of the progenitor cells. Furthermore, we show that disruption of viable ganglion formation leads to pathfinding errors of branchial motoneurons. Finally, our results suggest that the Eya-Six regulatory hierarchy also operates in the epibranchial placodal development. These findings uncover an essential function for Eya1 and Six1 as critical determination factors in acquiring both neuronal fate and neuronal subtype identity from epibranchial placodal progenitors. These analyses define a specific role for both genes in early differentiation and survival of the placodally derived cranial sensory neurons.

Animals↗
PubMed2000

Identification of synergistic signals initiating inner ear development.

Tissue manipulation experiments in amphibians more than 50 years ago showed that induction of the inner ear requires two signals: a mesodermal signal followed by a neural signal. However, the molecules mediating this process have remained elusive. We present evidence for mesodermal initiation of otic development in higher vertebrates and show that the mesoderm can direct terminal differentiation of the inner ear in rostral ectoderm. Furthermore, we demonstrate the synergistic interactions of the extracellular polypeptide ligands FGF-19 and Wnt-8c as mediators of mesodermal and neural signals, respectively, initiating inner ear development.

Animals↗
PubMed1996

[Temporal bone pathology in neonates with severe visceral anomalies].

It is well known that hearing loss is often associated with anomalads, syndromes involving multiple anomalies. The incidence is especially high in severe cases of facial and visceral malformations. However, otologic features remain unclear in patients with a sequence of various anomalies which can not be classified into any known syndrome. We examined 11 temporal bones from 6 patients with severe visceral anomalies, which could not be classified into any known systemic bone diseases, chromosomal abnormalities, or congenital metabolic disorders. Temporal bone pathology was compared with external and visceral anomalies in each case. The temporal bones had been removed at autopsy, fixed in 10% formaldehyde, decalcified and embedded in celloidin. Serial horizontal sections were made at 20 microns and every tenth section was stained with hematoxylin eosin. Most abnormalities in the middle and inner ear were found to have an ectodermal or mesodermal origin. Inner ear abnormalities were noted in 6 temporal bones from 3 patients; the predominant feature was hypoplasia of the semicircular canals. Middle ear abnormalities excluding residual mesenchymal tissue were noted in 5 temporal bones from 4 patients; the predominant feature was an abnormal course of the facial nerve. It was also suspected that auricular and maxillomandibular abnormalities, which are often associated with severe visceral anomalies, indicate a high incidence of disorders affecting the auditory and vestibular systems.

Abnormalities, Multiple↗
PubMed1984

Development of the apical ectodermal ridge in the chick wing bud.

Histological examination of the stage-18 to stage-23 chick wing bud apex revealed the following. Initially, the wing bud was covered by a cuboidal to columnar epithelium with an overlying periderm. Thickening of the apical ectoderm was not obvious until late stage 18 (36 pairs of somites), after the appearance of the wing bud. At late stage 18, cells of the inner layer of ectoderm had elongated slightly along an axis perpendicular to the epithelial-mesenchymal interface. Well-defined apical ectodermal ridge morphology, i.e., pseudostratified columnar epithelium with an overlying periderm, was not apparent until stage 20. Subsequently the ridge lengthened along the anteroposterior perimeter of the wing bud. We demonstrated histologically that the apical ectodermal ridge of the wing bud was asymmetric with respect to the anteroposterior axis, in that there was more ridge associated with posterior mesoderm. Other observations include the spatial and temporal location of a groove in the base of the thickest part of the ridge. The groove can be correlated with the specification of distal wing elements. The groove was first seen at stage 20 and became more prominent through stage 23. An anteroposterior progression of ectodermal cell death was also observed. This began at late stage 18 and continued through each of the stages examined.

Animals↗
PubMed1982

DNA content, mitotic activity, and incorporation of tritiated thymidine in the developing inner ear of the rat.

The rat inner ear is ectodermally derived from a region adjacent to the developing hindbrain. Beginning on day 8 of a 22-day gestational period, This zone of ectoderm first forms the otic placode, then the otocyst, and ultimately the definitive membranous labyrinth. This report provides an estimation of total DNA content of the developing inner ear, and hence an estimation of the total number of cells that comprise the inner ear at each developmental stage. The incorporation of 3H-thymidine indicates that most cells of the inner ear undergo DNA synthetic activity during gestational days 13 to 15. Radioautographic observations indicate a zone of DNA synthetic activity at the base of the outpocketing cochlear duct during early development. At the later stages of development, DNA synthesis is restricted to the cristae ampullares of the semicircular canals and the maculae of the utricle and the saccule. In contradistinction to the findings of other investigators, the statoacoustic ganglion complex undergoes terminal mitosis during gestational days 17 and 18. The gestational period between days 13 and 15 may prove to be a critical stage in normal otic development. The normal values of total DNA content and the number of cells that comprise the inner ear during development, established by these methods, can be compared with pathologic inner ears to provide quantitative means of assessing the damage in malformed inner ears. These values also form the baseline for future experimental studies of inner ear development.

Animals↗
PubMed2000

Development. Hear, hear, for the inner ear.

Although the development of the inner ear has been a favorite subject for biologists to study, it is not yet clear exactly which molecules are involved in the induction of the otic placode, the plug of embryonic ectoderm that will become the inner ear. In his Perspective, Graham takes us on an inner ear odyssey, explaining how the signaling molecules FGF-19 and Wnt-8c cooperate to induce formation of the otic placode (Ladher et al.).

Animals↗
PubMed2006

Radial intercalation of ciliated cells during Xenopus skin development.

Cells with motile cilia cover the skin of Xenopus tadpoles in a characteristic spacing pattern. This pattern arises during early development when cells within the inner layer of ectoderm are selected out by Notch to form ciliated cell precursors (CCPs) that then radially intercalate into the outer epithelial cell layer to form ciliated cells. When Notch is inhibited and CCPs are overproduced, radial intercalation becomes limiting and the spacing of ciliated cells is maintained. To determine why this is the case, we used confocal microscopy to image intercalating cells labeled using transplantation and a transgenic approach that labels CCPs with green fluorescent protein (GFP). Our results indicate that inner cells intercalate by first wedging between the basal surface of the outer epithelium but only insert apically at the vertices where multiple outer cells make contact. When overproduced, more CCPs are able to wedge basally, but apical insertion becomes limiting. We propose that limitations imposed by the outer layer, along with restrictions on the apical insertion of CCPs, determine their pattern of radial intercalation.

Animals↗
PubMed1995

Axial rotation in rat embryos: morphological analysis and microsurgical study on the role of the allantois.

In mouse and rat embryos, the embryonic disc develops within a cup-shaped "egg cylinder" and consists of an inner layer of ectoderm and an outer layer of endoderm. Because of this configuration, the embryo first develops in a dorsally flexed position and then undergoes "axial rotation" to a ventrally flexed position. In the present study, we first analyzed the morphological process of axial rotation in rat embryos using novel reference axes set in the egg cylinder that remained invariant during the process. Our new perspective allowed us to demonstrate that the process consists of three movements which start at different stages of development: twisting of the upper body at stage 12/s7-8, twisting of the middle body at stage 13/s11-12, and twisting of the lower body (so called "tail") at stage 14/s15-16. Axial rotation is an interesting developmental event not only because it is such a dynamic process but also because it is one of the earliest morphological signs of body asymmetry. This asymmetry is strongly biased in that the tail almost always finishes up on the right side of the embryo for reasons that are still unknown. In the second part of the study, we performed microsurgical experiments to extend our previous finding that removal of the allantois results in random determination of tail sidedness. We demonstrated that an allantois transplanted from another embryo can prevent this abnormal sidedness in an embryos whose allantois had been removed and that transecting the allantois did not lead to abnormal tail sidedness. A possible explanation is that the allantois produces a chemical factor that controls tail sidedness.

Allantois↗
PubMed1983

Fibronectin in early amphibian embryos. Migrating mesodermal cells contact fibronectin established prior to gastrulation.

The three-dimensional organisation of fibronectin (FN) in early amphibian embryos (Ambystoma mexicanum, Pleurodeles waltlii) was studied with the use of antibodies against amphibian-FN. Immunofluorescence labelling was performed on whole-mount specimens. It was shown that before gastrulation an extensive extracellular network consisting of anastomosed FN-fibrils underlies the roof of the blastocoel prior to the migration of mesodermal cells. Initially, FN-fibrils develop radially on the inner surface of ectodermal cells (early blastula stage) and become confluent there to elaborate a fibrillar network. During the late blastula stage the entire surface of the blastocoel roof (presumptive ectoderm and mesoderm) is covered by the FN-rich extracellular matrix. As gastrulation proceeds, the migrating mesodermal cells interact with the FN-fibrillar network. The results suggest that the FN-containing extracellular matrix plays an important role in mediating mesodermal cell-substratum contacts in gastrulating embryos. No FN-fibrils were found in other parts of the embryo.

Ambystoma↗
PubMed1983

Differential expression of lectin receptors in germ layers of the mouse egg cylinder and teratocarcinomas.

Receptors for three lectins with restricted specificities, namely fucose-binding protein of Lotus tetragonolobus (FBP), peanut agglutinin (PNA) and Dolichos biflorus agglutinin (DBA), were distinctively located in 6- and 7-day mouse embryos and in embryoid bodies of teratocarcinoma OTT6050 grown in vivo. Thus, FBP reacted mainly with the inner cells (embryonic ectoderm and teratocarcinoma stem cells), DBA reacted with the outer cells (endoderm) and PNA reacted with all the germ layers including mesoderm. Upon in vitro culture of the embryoid bodies, the exposed stem cells express DBA receptors. Since the receptors for the three lectins in teratocarcinomas are known to be carried by the large carbohydrate chains characteristic of early embryonic cells, the present result suggests that terminal structure of the large carbohydrates is altered according to the direction of the differentiation or to the position of the cells in embryos and in teratocarcinomas.

Amnion↗
PubMed1999

PAX6 expression in the developing human eye.

AIMS: To investigate the changes in PAX6 expression in the developing human eye. METHODS: Six developing human eyes from 6 to 22 weeks' gestation were evaluated. Frozen sections were immunohistochemically stained with monoclonal antibody to chick Pax6 (amino acids 1-223). To verify antibody specificity, western blot analysis was carried out using cell lysates from P19 cells transfected with the human PAX6 gene. RESULTS: Western blot analysis demonstrated that the antibody reacted to human PAX6 protein. Positive immunostainings for PAX6 were seen in the surface ectoderm, lens vesicle, inner and outer layers of the optic cup, and optic stalk at 6 weeks, and in the corneal epithelia and conjunctiva, lens, and non-pigmented ciliary epithelia from 8 to 22 weeks. In the retina, positive cells were seen in the entire retina from 8 to 10 weeks, and were restricted to the ganglion cell layer and the inner and outer portions of the inner nuclear layer after 21 weeks. CONCLUSIONS: PAX6 is expressed on the surface and neuroectoderms at an early stage, then in the differentiating cells in the cornea, lens, ciliary body, and retina through development. PAX6 may play a role in determining cell fate in the morphogenesis of various human ocular tissue.

Antibody Specificity↗
PubMed1997

Murine gastrulation requires HNF-4 regulated gene expression in the visceral endoderm: tetraploid rescue of Hnf-4(-/-) embryos.

Immediately prior to gastrulation the murine embryo consists of an outer layer of visceral endoderm (VE) and an inner layer of ectoderm. Differentiation and migration of the ectoderm then occurs to produce the three germ layers (ectoderm, embryonic endoderm and mesoderm) from which the fetus is derived. An indication that the VE might have a critical role in this process emerged from studies of Hnf-4(-/-) mouse embryos which fail to undergo normal gastrulation. Since expression of the transcription factor HNF-4 is restricted to the VE during this phase of development, we proposed that HNF-4-regulated gene expression in the VE creates an environment capable of supporting gastrulation. To address this directly we have exploited the versatility of embryonic stem (ES) cells which are amenable to genetic manipulation and can be induced to form VE in vitro. Moreover, embryos derived solely from ES cells can be generated by aggregation with tetraploid morulae. Using Hnf-4(-/-) ES cells we demonstrate that HNF-4 is a key regulator of tissue-specific gene expression in the VE, required for normal expression of secreted factors including alphafetoprotein, apolipoproteins, transthyretin, retinol binding protein, and transferrin. Furthermore, specific complementation of Hnf-4(-/-) embryos with tetraploid-derived Hnf-4(+/+) VE rescues their early developmental arrest, showing conclusively that a functional VE is mandatory for gastrulation.

Animals↗
PubMed2006

Wnt signals mediate a fate decision between otic placode and epidermis.

The otic placode, the anlagen of the inner ear, develops from an ectodermal field characterized by expression of the transcription factor Pax2. Previous fate mapping studies suggest that these Pax2(+) cells will give rise to both otic placode tissue and epidermis, but the signals that divide the Pax2(+) field into placodal and epidermal territories are unknown. We report that Wnt signaling is normally activated in a subset of Pax2(+) cells, and that conditional inactivation of beta-catenin in these cells causes an expansion of epidermal markers at the expense of the otic placode. Conversely, conditional activation of beta-catenin in Pax2(+) cells causes an expansion of the otic placode at the expense of epidermis, and the resulting otic tissue expresses exclusively dorsal otocyst markers. Together, these results suggest that Wnt signaling acts instructively to direct Pax2(+) cells to an otic placodal, rather than an epidermal, fate and promotes dorsal cell identities in the otocyst.

Animals↗
PubMed1983

Comparative study of extracellular fibrils on the ectodermal layer in gastrulae of five amphibian species.

Previous studies have shown the presence of a network of extracellular fibrils on the inner surface of the ectodermal layer of the Ambystoma maculatum gastrulae. The alignment of the network along the blastopore-animal pole axis has suggested that the network of fibrils guides the migrating mesodermal cells in gastrulae by contact guidance. We have also shown that these fibrils can be deposited on substrata by explanted embryonic fragments and that substrata conditioned in this manner support directed cell migration. In this study, we found that the appearance of the fibrils in the embryos coincides with the start of cell migration towards the animal pole. Gastrulae of three urodele species examined (A. maculatum, A. mexicanum and Cynops pyrrhogaster) have similar dense networks of fibrils. Xenopus laevis gastrulae also have similar fibrils but fewer fibrils compared to urodele embryos. Rana pipiens gastrulae have very few extracellular fibrils. The scarcity of the fibrils in anuran species may be related to the differences in arrangement of mesodermal cells during migration.

Ambystoma↗
PubMed2000

Competence, specification and commitment in otic placode induction.

The inner ear is induced from cranial ectoderm adjacent to the hindbrain. Despite almost a century of study, the molecular mechanisms of inner ear induction remain obscure. We have identified four genes expressed very early in the anlage of the inner ear, the otic placode. Pax-2, Sox-3, BMP-7 and Notch are all expressed in placodal ectoderm from the 4-5 somite stage (ss) onwards, well before the otic placode becomes morphologically visible at the 12-14ss. We have used these four molecular markers to show that cranial ectoderm becomes specified to form the otic placode at the 4-6ss, and that this ectoderm is committed to a placodal fate by the 10ss. We also demonstrate that much of the embryonic ectoderm is competent to generate an otic placode if taken at a sufficiently early age. We have mapped the location of otic placode-inducing activity along the rostrocaudal axis of the embryo, and have determined that this activity persists at least until the 10ss. Use of the four molecular otic placode markers suggests that induction of the otic placode in birds occurs earlier than previously thought, and proceeds in a series of steps that are independently regulated.

Animals↗