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Volker Schmid

Publications and source records attributed to Volker Schmid.

16 recordsLinked to original sources

Animal pole determinants define oral-aboral axis polarity and endodermal cell-fate in hydrozoan jellyfish Podocoryne carnea.

Cnidarians, in contrast with bilaterians, are generally considered to exhibit radial symmetry around a single body axis (oral-aboral) throughout their life-cycles. We have investigated how the oral-aboral axis is established in the hydrozoan jellyfish Podocoryne carnea. Vital labeling experiments showed that the oral end of the blastula derives from the animal pole region of the egg as has been demonstrated for other cnidarian species. Gastrulation is restricted to the oral pole such that the oral 20% of blastula cells give rise to endoderm. Unexpectedly, bisection experiments at the 8-cell stage showed that animal regions are able to develop into normally polarized larvae, but that vegetal (aboral) blastomeres completely fail to develop endoderm or to elongate. These vegetal-derived larvae also failed to polarize, as indicated by a lack of oral-specific RFamide-positive nerve cells and a disorganized tyrosinated tubulin-positive nerve net. A different result was obtained following bisection of the late blastula stage: aboral halves still lacked the capacity to develop endoderm but retained features of axial polarity including elongation of the larva and directional swimming. These results demonstrate for the first time in a cnidarian the presence of localized determinants responsible for axis determination and endoderm formation at the animal pole of the egg. They also show that axial polarity and endoderm formation are controlled by separable pathways after the blastula stage.

Alkaline Phosphatase↗

A two-component model for counts of infectious diseases.

We propose a stochastic model for the analysis of time series of disease counts as collected in typical surveillance systems on notifiable infectious diseases. The model is based on a Poisson or negative binomial observation model with two components: a parameter-driven component relates the disease incidence to latent parameters describing endemic seasonal patterns, which are typical for infectious disease surveillance data. An observation-driven or epidemic component is modeled with an autoregression on the number of cases at the previous time points. The autoregressive parameter is allowed to change over time according to a Bayesian changepoint model with unknown number of changepoints. Parameter estimates are obtained through the Bayesian model averaging using Markov chain Monte Carlo techniques. We illustrate our approach through analysis of simulated data and real notification data obtained from the German infectious disease surveillance system, administered by the Robert Koch Institute in Berlin. Software to fit the proposed model can be obtained from http://www.statistik.lmu.de/ approximately mhofmann/twins.

Algorithms↗

BMP2/4 and BMP5-8 in jellyfish development and transdifferentiation.

Bone morphogenetic proteins (BMPs) have key roles in gastrulation, mesoderm induction and axial patterning. The multitude of bilaterian BMPs employed in these morphogenetic processes contrasts starkly with the scarcity of BMPs in Cnidaria, the most basal eumetazoan phylum. In coral, sea anemone and hydra species, BMPs have been found to be associated with larval and polyp axial patterning. In the hydrozoan jellyfish Podocoryne (Hydractinia) carnea the BMP2/4 and BMP5-8 genes are expressed unilaterally in the larva, corroborating a possible role in larval axial development. With the focal area of BMP expression in the anterior region, however, the jellyfish larva may have a developmental reversal of spatial polarity compared to the anthozoan larva. In medusa development, BMP genes are expressed in divergent expression territories within the presumptive radial canals and in various parts of the endoderm, indicative of an involvement in mesoderm patterning and gastrovascular system formation reminiscent of bilaterian BMP functions. In addition, the BMP2/4 and BMP5-8 genes may play roles in wound response and dedifferentiation or S-phase re-entry, respectively, as the former is expressed in striated muscle cells immediately after excision from the bell and the latter in the initial phase of muscle cell transdifferentiation.

Amino Acid Sequence↗

Mesodermal anatomies in cnidarian polyps and medusae.

The cellular and developmental analysis of evolutionary-conserved genes directing bilaterian mesodermal and myogenic cell fate previously identified the hydromedusan entocodon and its differentiation product, the striated muscle, as mesodermal derivatives. In view of these findings we presented a hypothesis disputing the diploblast classification of cnidarians without providing further explanations for the apparent diploblasty of the polyp stage and the formation of the subepidermal striated muscle in those Medusozoa lacking the entocodon nodule (Seipel and Schmid, 2005). Hence we carried out a systematic review of the histological and experimental evidence for mesodermal differentiations in cnidarians. In anthozoan and scyphozoan but not in hydrozoan polyps the presumptive mesodermal elements include amoeboid cells, the mesentery retractor muscles and scleroblasts, all of which are embedded or deeply rooted in the extracellular matrix (mesoglea) and derive from the ectoblastemal cells invading the extracellular matrix from the gastrulation site during or shortly after endoderm formation. These data lend further support to the cnidarian mesodermate hypothesis, whereby cnidarians and bilaterians share a common triploblast ancestor, the Urtriploblast, a small, motile, possibly medusa-like organism that did not feature a sessile polyp stage in its life cycle. As a consequence the diploblasty of the hydrozoan polyps may represent a derived morphology resulting from heterochronic modulations of the gastrulation process after endoderm formation.

Animals↗

Evolution of striated muscle: jellyfish and the origin of triploblasty.

The larval and polyp stages of extant Cnidaria are bi-layered with an absence of mesoderm and its differentiation products. This anatomy originally prompted the diploblast classification of the cnidarian phylum. The medusa stage, or jellyfish, however, has a more complex anatomy characterized by a swimming bell with a well-developed striated muscle layer. Based on developmental histology of the hydrozoan medusa this muscle derives from the entocodon, a mesoderm-like third cell layer established at the onset of medusa formation. According to recent molecular studies cnidarian homologs to bilaterian mesoderm and myogenic regulators are expressed in the larval and polyp stages as well as in the entocodon and derived striated muscle. Moreover striated and smooth muscle cells may have evolved directly and independently from non-muscle cells as indicated by phylogenetic analysis of myosin heavy chain genes (MHC class II). To accommodate all evidences we propose that striated muscle-based locomotion coevolved with the nervous and digestive systems in a basic metazoan Bauplan from which the ancestors of the Ctenophora (comb jellyfish), Cnidaria (jellyfish and polyps), as well as the Bilateria are derived. We argue for a motile tri-layered cnidarian ancestor and a monophyletic descent of striated muscle in Cnidaria and Bilateria. As a consequence, diploblasty evolved secondarily in cnidarian larvae and polyps.

Animals↗

The Sine oculis/Six class family of homeobox genes in jellyfish with and without eyes: development and eye regeneration.

The development of visual organs is regulated in Bilateria by a network of genes where members of the Six and Pax gene families play a central role. To investigate the molecular aspects of eye evolution, we analyzed the structure and expression patterns of cognate members of the Six family genes in jellyfish (Cnidaria, Hydrozoa), representatives of a basal, non-bilaterian phylum where complex lens eyes with spherical lens, an epidermal cornea, and a retina appear for the first time in evolution. In the jellyfish Cladonema radiatum, a species with well-developed lens eyes in the tentacle bulbs, Six1/2-Cr and Six3/6-Cr, are expressed in the eye cup. Six4/5-Cr is mainly expressed in the manubrium, the feeding, and sex organ. All three Six genes are expressed in different subsets of epidermal nerve cells, possibly of the RFamide type which are part of a net connecting the different eyes with each other and the effector organs. Furthermore, expression is found in other tissues, notably in the striated muscle. During eye regeneration, expression of Six1/2-Cr and Six3/6-Cr is upregulated, but not of Six4/5-Cr. In Podocoryne carnea, a jellyfish without eyes, Six1/2-Pc and Six3/6-Pc are also expressed in the tentacle bulbs, Six1/2-Pc additionally in the manubrium and striated muscle, and Six3/6-Pc in the mechanosensory nematocytes of the tentacle. The conserved gene structure and expression patterns of all Cladonema Six genes suggest broad conservation of upstream regulatory mechanisms in eye development.

Animals↗

Developmental and evolutionary aspects of the basic helix-loop-helix transcription factors Atonal-like 1 and Achaete-scute homolog 2 in the jellyfish.

The close functional link of nerve and muscle cells in neuromuscular units has led to the hypothesis of a common evolutionary origin of both cell types. Jellyfish are well suited to evaluate this theory since they represent the most basal extant organisms featuring both striated muscle and a nervous system. Here we describe the structure and expression of two novel genes for basic helix-loop-helix (bHLH) transcription factors, the Achaete-scute B family member Ash2 and the Atonal-like gene Atl1, in the hydrozoan jellyfish Podocoryne carnea. Ash2 is expressed exclusively in larval and adult endoderm cells and may be involved in differentiation of secretory cells. Atl1 expression is more widespread and includes the developing striated muscle as well as mechanosensory and nerve cell precursors in the medusa tentacles. Moreover, Atl1 expression is upregulated in proliferating nerve cell precursors arising from adult striated muscle cells by transdifferentiation in vitro. Likewise, the neuronal marker gene NP coding for the RFamide neuropeptide is expressed not only in mature nerve cells but also transiently in the developing muscle. The molecular evidence is concurrent to the hypothesis that muscle and nerve cells are closely linked in evolution and derive from a common myoepithelial precursor.

Amino Acid Sequence↗

Basic leucine zipper transcription factors C/EBP and MafL in the hydrozoan jellyfish Podocoryne carnea.

Members of the CCAAT/enhancer binding protein (C/EBP) and the Maf protein subfamilies have been characterized in a variety of bilaterian organisms. This is the first report of C/EBP and MafL genes in a basal organism, the hydrozoan jellyfish Podocoryne carnea. Transcripts of both genes are present in all life cycle stages: egg, embryo, larva, polyp, and medusa. During early development, both factors appear to regulate metamorphosis of the larva to the primary polyp. Both genes are also expressed in the striated muscle of the developing and adult medusa. During in vitro transdifferentiation of striated muscle cells to smooth muscle and nerve cells, C/EBP is continuously expressed, whereas MafL expression is turned off during transdifferentiation and reactivated when nerve cells differentiate. Thus, both factors may be involved in muscle and nerve cell differentiation. In the mature medusa both genes are also implicated in gametogenesis. Developmental and evolutionary aspects of the gene structures and expression patterns are discussed.

Amino Acid Sequence↗

Homologs of vascular endothelial growth factor and receptor, VEGF and VEGFR, in the jellyfish Podocoryne carnea.

Vascular endothelial growth factors (VEGF) are the major inducers of vasculogenesis and angiogenesis in vertebrates. Their effects are mediated by receptor tyrosine kinases of the VEGF receptor (VEGFR) family located on endothelial cells and include stimulation of cell survival, proliferation, migration, and tube formation as well as regulation of vascular permeability. Here, we report the presence of VEGF and VEGFR homologous genes in a basal invertebrate of the phylum Cnidaria. The marine jellyfish Podocoryne carnea features a gastrovascular system consisting of the feeding organ, or manubrium, the radial and ring canals, and the tentacle bulbs. Expression analysis indicates that both genes are involved in tentacle and gastrovascular canal formation, indicating an early recruitment of the VEGF signalling pathway for morphogenetic processes leading to tube formation in metazoans. The evolutionary origin of the VEGF signalling pathway resides in the common ancestor of the Cnidaria and Bilateria.

Amino Acid Sequence↗

Bayesian extrapolation of space-time trends in cancer registry data.

We apply a full Bayesian model framework to a dataset on stomach cancer mortality in West Germany. The data are stratified by age group, year, and district. Using an age-period-cohort model with an additional spatial component, our goal is to investigate whether there is evidence for space-time interactions in these data. Furthermore, we will determine whether a period-space or a cohort-space interaction model is more appropriate to predict future mortality rates. The setup will be fully Bayesian based on a series of Gaussian Markov random field priors for each of the components. Statistical inference is based on efficient algorithms to block update Gaussian Markov random fields, which have recently been proposed in the literature.

Bayes Theorem↗

The germ line and somatic stem cell gene Cniwi in the jellyfish Podocoryne carnea.

In most animal phyla from insects to mammals, there is a clear division of somatic and germ line cells. This is however not the case in plants and some animal phyla including tunicates, flatworms and the basal phylum Cnidaria, where germ stem cells arise de novo from somatic cells. Piwi-like genes represent essential stem cell genes in diverse multicellular organisms. The cnidarian Piwihomolog Cniwiwas cloned from Podocoryne carnea, a hydrozoan with a full life cycle. CniwiRNA is present in all developmental stages with highest levels in the egg and the medusa. In the adult medusa, Cniwi expression is prominent in the gonads where it likely functions as a germ stem cell gene. The gene is also expressed, albeit at low levels, in differentiated somatic cells like the striated muscle of the medusa. Isolated striated muscle cells can be induced to transdifferentiate into smooth muscle cells which proliferate and differentiate into nerve cells. Cniwi expression is upregulated transiently after induction of transdifferentiation and again when the emerging smooth muscle cells proliferate and differentiate. The continuous low-level expression of an inducible stem cell gene in differentiated somatic cells may underlie the ability to form medusa buds from polyp cells and explain the extraordinary transdifferentation and regeneration potential of Podocoryne carnea.

Aging↗

Evolutionary aspects of developmentally regulated helix-loop-helix transcription factors in striated muscle of jellyfish.

The function of basic helix-loop-helix (bHLH) proteins in cell differentiation was shown to be conserved from Drosophila to vertebrates, exemplified by the function of MyoD in striated muscle differentiation. In phylogeny striated muscle tissue appears first in jellyfish and the question of its evolutionary position is controversially discussed. For this reason we have studied the developmental role of myogenic bHLH genes in medusa development. Based on their dimerization ability, four genes of the bHLH family of transcription factors were isolated from the hydrozoan jellyfish Podocoryne carnea. While the proteins Id and Ash group with cognate family members from bilaterians, Net-like and JellyD1 could not be unequivocally classified. Id is expressed during the medusa budding process and in the adult medusa, Ash and Net-like are expressed in all life cycle stages from egg to adult medusa and JellyD1 is expressed in the blastula and gastrula stages, the planula larva, and in late medusa bud stages. The dimerization specificity, the expression pattern, and the conservation of two residues specific for a MyoD bHLH domain suggest that JellyD1 is related to an ancestral MyoD gene. Id, Net-like, and JellyD1 are either expressed in the entocodon or its derived tissues, the striated and smooth muscle of the bell. These findings strengthen the hypothesis that the entocodon is a mesoderm-like structure and that the common ancestor of Cnidaria and Bilateria was more complex in cell-type architecture and body organization than commonly thought.

Amino Acid Sequence↗

Conservation of Brachyury, Mef2, and Snail in the myogenic lineage of jellyfish: a connection to the mesoderm of bilateria.

One major difference between simple metazoans such as cnidarians and all the bilaterian animals is thought to involve the invention of mesoderm. The terms diploblasts and triploblasts are therefore, often used to group prebilaterian and bilaterian animals, respectively. However, jellyfish contain well developed striated and smooth muscle tissues that derive from the entocodon, a mesoderm-like tissue formed during medusa development. We investigated the hypothesis, that the entocodon could be homologous to the third germ layer of bilaterians by analyzing the structures and expression patterns of the homologues of Brachyury, Mef2, and Snail in the jellyfish Podocoryne carnea. These are regulatory genes from the T-box, MADS-box and zinc finger families known to play important roles in bilaterian mesoderm patterning and muscle differentiation. The sequence and expression data demonstrate that the genes are structurally and functionally conserved and even more similar to humans or other deuterostomes than to protostome model organisms such as Drosophila or Caenorhabditis elegans. Based on these data we conclude that the common ancestor of the cnidarians and bilaterians not only shared genes that play a role in regulating myogenesis but already used them to develop and differentiate muscle systems similar to those of triploblasts.

Animals↗

Evolutionary conservation of the chromatin modulator Polycomb in the jellyfish Podocoryne carnea.

Polycomb-group (PcG) proteins form chromatin-associated multimeric complexes, which are responsible for the maintenance of the transcriptionally repressive state of regulatory genes during development. We have isolated a Polycomb homologue of the hydrozoan Podocoryne carnea by a PCR-based approach. Our results demonstrate that structure and function of Polycomb-group proteins have been conserved in evolution from cnidarians to vertebrates since Podocoryne Polycomb interacts in yeast with mouse dinG/RING1B, an interaction partner of the mouse Polycomb homologue MPc3. Polycomb is expressed throughout the life cycle of Podocoryne. In situ hybridization reveals a differential expression pattern in proliferating and differentiating tissues of the developing medusa bud. In the transdifferentiation of activated isolated striated muscle of the medusa to smooth muscle and RFamide-positive nerve cells, Polycomb expression is strongly increased when differentiation into nerve cells occurs.

Amino Acid Sequence↗

Cnidarians as a model system for understanding evolution and regeneration.

Hydra and Podocolyne are two cnidarian animals which provide complementary advantages for analysing developmental mechanisms possibly reflecting the basic developmental processes shared by most bilaterians. Interestingly, these mechanisms remain accessible all along the life of these animals, which bud and regenerate, whatever their age. The Hydra polyp permits a direct study of the molecular cascades linking amputation to regeneration. Podocoryne displays a complete life cycle, polyp and medusa stages with a fast and inducible sexual cycle and an unparalleled In vitro transdifferentiation potential. In both cases, a large number of evolutionarily conserved molecular markers are available, and analysis of their regulation highlights the molecular mechanisms which underly pattern formation in these two species.

Animals↗

nanos expression at the embryonic posterior pole and the medusa phase in the hydrozoan Podocoryne carnea.

Summary The distinction between soma and germline is an important process in the development of animals with sexual reproduction. It is regulated by a number of germline-specific genes, most of which appear conserved in evolution and therefore can be used to study the formation of the germline in diverged animal groups. Here we report the isolation of two orthologs of one such gene, nanos (nos), in the cnidarian Podocoryne carnea, a species with representative zoological features among the hydrozoans. By studying nos gene expression throughout the Podocoryne biphasic life cycle, we find that the germline differentiates exclusively during medusa development, whereas the polyp does not contribute to the process. An early widespread nos expression in developing medusae progressively refines into a mainly germline-specific pattern at terminal stages of medusa formation. Thus, the distinction between germline and soma is a late event in hydrozoan development. Also, we show that the formation of the medusa is a de novo process that relies on active local cell proliferation and differentiation of novel cell and tissue types not present in the polyp, including nos-expressing cells. Finally, we find nos expression at the posterior pole of Podocoryne developing embryos, not related to germline formation. This second aspect of nos expression is also found in Drosophila, where nos functions as a posterior determinant essential for the formation of the fly abdomen. This raises the possibility that nos embryonic expression could play a role in establishing axial polarity in cnidarians.

Amino Acid Sequence↗