Academic literature on the topic 'Hemichordata'

Here, we report the discovery and characterization of biominerals in the acorn worms Saccoglossus bromophenolosus and Ptychodera flava galapagos (Phylum: Hemichordata). Using electron microscopy, X-ray microprobe analyses and confocal Raman spectroscopy, we show that hemichordate biominerals are small CaCO 3 aragonitic elements restricted to specialized epidermal structures, and in S. bromophenolosus, are apparently secreted by sclerocytes. Investigation of urchin biomineralizing proteins in the translated genome and expressed sequence tag (EST) libraries of Saccoglossus kowalevskii indicates that three members of the urchin MSP-130 family, a carbonic anhydrase and a matrix metaloprotease are present and transcribed during the development of S. kowalevskii . The SM family of proteins is absent from the hemichordate genome. These results increase the number of phyla known to biomineralize and suggest that some of the gene-regulatory ‘toolkit’, if not mineralized tissue themselves, may have been present in the common ancestor to hemichordates and echinoderms.

AbstractYuknessia Walcott, 1919 recently was transferred from the green algae to the Phylum Hemichordata on the basis of new details observed for the type species, Y. simplex, from the Burgess Shale Formation (Cambrian Stage 5) of British Columbia. This has prompted reexamination of material attributed to Yuknessia from various Cambrian localities in South China. Findings preclude both a Yuknessia and a hemichordate affinity for all of the Chinese study material, and most of this material is formally transferred to Fuxianospira Chen and Zhou, 1997, a taxon common in the Chengjiang biota. Comparable material from the Cambrian Marjum, Wheeler, and Burgess Shale formations of North America is also assigned to Fuxianospira, and this reassignment expands both the paleogeographic and stratigraphic range of this taxon. All aspects of the study specimens, including details obtained from scanning electron microscopy, are consistent with an algal affinity, as proposed in the original descriptions of the Chinese material.

The work embodied in this thesis concerns the host defence mechanisms of the hemichordate, <i>Saccoglossus ruber</i> and the cephalochordate, <i>Branchiostoma lanceolatum</i>. Initially the cellular repsonses of these animals towards experimentally introduced foreign particles were examined. <i>S. ruber</i> possesses avidly phagocytic coelomocytes which were studied <i>in vitro</i> and <i>in vivo</i>, and shown by light and electron microscopy to be of a single morphological type. These coelomocytes probably originate from the coelomic lining, but autoradiography failed to substantiate this. The cellular response in <i>B. lanceolatum</i> towards wounding and subcutaneously injected bacteria was very limited. Even after 7 days, a relatively small number of coelomocytes accumulated at the site of injection. These coelomocytes contained phagocytosed bacteria and formed the beginning of nodule-like structures. In the majority of sections, the bacteria were observed surrounded by an unidentifiable amorphous material. Haemagglutinins and antibacterial factors were also detected in whole body homogenates of <i>S. ruber</i> and <i>B. lanceolatum</i>. The haemagglutinins from both animals were proteinaceous and functioned independently of Ca<SUP>++</SUP> and Mg<SUP>++</SUP>. The <i>S. ruber</i> agglutinin showed specificity for D-galactose containing carbohydrates, whereas no clear-cut carbohydrate specificity could be discerned with the <i>B. lanceolatum</i> agglutinin. The <i>S. ruber</i> agglutinin was also associated with the mucus covering the body surface, and the <i>B. lanceolatum</i> agglutinin with mucus in the pharynx. Antibacterial activity in <i>S. ruber</i> was strongest against natural bacterial isolates. The active factor, which may be a bromophenol, was heat stable, dialyzable, bactericidal in its mode of action and also associated with the body surface mucus. The <i>B. lanceolatum</i> antibacterial factor was bacteriostatic, probably proteinaceous, heat labile and operated over a wide temperature range. At a protein concentration above 5 mg ml<SUP>-1</SUP>, the antibacterial proteins aggregated and could be disassociated only at high pH. A role for these humoral factors in the defence systems of the experimental animals is discussed.

The aim of this research is to gain new insights into the evolution of deuterostomes through molecular developmental investigation of pterobranchs. This thesis consists of three major parts. In the first part, I investigated the seasonal reproductive activity of the pterobranch species Rhabdoplellra compacta. By analyzing large numbers of specimens for two years, I found that their reproductive activity has a peak in June. In the second part, I investigated the origin of vertebrate left-right axis patterning using anatomical as well as molecular studies on R. compacta. I found that R. compacta indeed have anatomical left-right asymmetry in the body. However, contrary to the traditional description, the direction was random, rather than directional as traditionally described. Together with further molecular analyses and previous morphological studies in enteropneusts, I suggest that precise left-right patterning already existed before the divergence of deuterostomes, but underwent degeneration during the evolution of the hemichordate lineage. The molecular analyses brought about two unexpected discoveries. One is that the expression patterns of the gene I examined were completely different from those of enteropneusts. Importantly, expression of the hedgehog gene in pterobranchs was found in the dorsal midline as in chordates, suggesting that the dorsal side of pterobranchs may be comparable to the dorsal side of chordates. The other is that there are unusual mutations in the autoproteolytic cleavage region of hedgehog proteins in the hemichordate lineage. In the third part, I analyzed whether these mutations alter efficiency of autoproteolytic cleavage and the signaling function, using Drosophila genetics. It was found that an enteropneust-type mutation decreases the efficiency of autoproteolytic cleavage whereas a pterobranch-type does not. Since this autoproteolytic cleavage is essential for the signaling function of hedgehog proteins, I suggest that hedgehog underwent unique functional evolution in the enteropneust lineage, which requires less efficiency of signaling function.

Le phylum Hemichordata est composé exclusivement d'organismes marins et, avec les embranchement Echinodermata et chordata, il forme le groupe des Deutérostomes sur l'arbre de la vie des animaux. Dans les chapitres d'introduction et le deuxième, je donne un aperçu des hémichordés, y compris les enteropneustes solitaires et les pterobranches coloniaux et je les défini dans un contexte évolutif ou phylogénétique. Les enteropneustes sont souvent considérés comme le meilleur proxy vivant de l'ancêtre des deutérostomes. Les ptérobranches comprennent les Cephalodiscida et les Graptolithina. Les graptolites (graptos = écrit, lithos = roche) sont principalement représentés par des espèces fossiles remontant à la Période Cambrienne, il y a plus de 500 millions d'années. Ces «écritures dans la roche» sont largement connues et étudiées par les paléontologues et sont si abondantes qu'elles sont utilisées comme fossiles indicateurs pour identifier les couches sédimentaires. Les graptolites sont éteints sauf pour cinq espèces benthiques appartenant au genre Rhabdopleura, membres de la famille Rhabdopleuridae, que j'examine en détail dans le chapitre trois. Rhabdopleura recondita de la mer Méditerranée fait l'objet de cette thèse. Il est courant le long des côtes sud de l'Italie d'où je l’ai échantillonné en plongée sous- marine. Il est inhabituel que des colonies résident cachées à l'intérieur de la zoaria des bryozoaires morts. Seuls les tubes érigés font saillie à partir de la matrice de l'hôte. Les chapitres quatre et cinq sont les contributions les plus significatives de cette thèse, avec un accent sur les tubes de R. recondita. Le chapitre quatre fournit des observations de la construction de tubes par R. recondita gardé en captivité. J'ai observé la capacité des larves, des zooïdes et des colonies à sécréter de nouveaux tubes en présence et en l'absence du matériel hôte du zoarium bryozoaire. Nous avons découvert que la colonisation larvaire et la sécrétion du dôme peuvent se produire sans l'hôte bryozoaire, mais la croissance continue de la colonie nécessite le substrat de l'hôte. Les zooïdes adultes ne peuvent reformer de nouveaux tubes que s'ils sont capables de s'abriter à l'intérieur du matériel hôte. Un résultat surprenant des observations des zooïdes a été la sécrétion d'un opercule et d'un tube évasé. Les colonies qui avaient des tubes érigés enlevés ont pu fabriquer de nouveaux tubes, mais à un faible nombre. Une étude parallèle a été réalisée sur des colonies dont les tubes avaient été retirés, puis cultivées dans des canaux à quatre vitesses d'écoulement. Cette expérience a été conçue pour induire une réponse plastique phénotypique à l'écoulement. Au lieu de cela, je n'ai trouvé aucune différence significative dans la longueur du tube ou le nombre de tubes en réponse à quatre vitesses d'écoulement. Ce résultat suggère que le développement du tube de R. recondita peut être canalisé ou fixé. Il est significatif car il suggère que de petites différences qui distinguent les espèces primitives de graptolites encroûtantes sont bonnes. Le chapitre cinq porte sur la composition des tubes de R. recondita. Plusieurs hypothèses et de nombreuses analyses ont été faites sur ce sujet, mais aucune n'a été concluante. J'utilise ici la génomique et la bioinformatique, l'immunochimie et la spectroscopie et rejette les hypothèses selon lesquelles les tubes sont de la kératine ou de la cellulose. Au lieu de cela, j'ai trouvé huit gènes de chitine synthase dans le génome et le trascriptome, un complexe composé d'un polysaccharide semblable à la chitine, d'une protéine, d'un acide gras et de composants élémentaires inattendus. Cette étude est significative car elle ferme la porte sur une ancienne hypothèse de composition de tube de graptolite et révèle qu'il s'agit d'une structure complexe comprenant de la chitine. Le chapitre de conclusion est un bref résumé des résultats et une réflexion sur les aveenues potentiellement fructueuse pour des recherches futures.<br>The phylum Hemichordata is comprised of exclusively marine organisms, and together with the Echinodermata and Chordata forms the Deuterostomia branch on the animal tree of life. In the introductory and second chapters I provide a background on Hemichordata including the solitary Enteropneusta and the colonial Pterobranchia and define them in an evolutionary or phylogenetic context. The enteropneusts are often regarded as the best living proxy of the deuterostome ancestor. Pterobranchs, include the Cephalodiscida and Graptolithina. Graptolites (graptos=written, lithos=rock) are mostly represented by fossil species dating back to the Cambrian Period, more than 500 million years ago. These “writings in the rock” are widely known and studied by paleontologists and are so abundant that they are used as index fossils to identify sedimentary layers. Graptolites are extinct but for five benthic species belonging to the genus Rhabdopleura, members of the Rhabdopleurida, which I extensively review in chapter three. Rhabdopleura recondita from the Mediterranean Sea is the subject of this thesis. It is common along the south coasts of Italy from where I sample it by SCUBA diving. It is unusual in that colonies reside hidden inside of the zoaria of dead bryozoans. Only erect tubes project from the host matrix. Chapters four and five are the most significant contributions of this thesis, with a focus on R. recondita tubes. Chapter four provides observations of tube building by R. recondita kept in captivity. I observed larvae, zooids and colonies abilities to secrete new tubes in the presence and absence of the bryozoan zoarium host material. We discovered that larval settlement and dome secretion can occur without the bryozoan host, but the continued growth of the colony requires the host substrate. Adult zooids can reform new tubes only if they are able to shelter inside of host material. A surprising result from the zooid observations was the secretion of an operculum and a flared tube. Colonies that had erect tubes removed were able to make new tubes, but fewer in number. A parallel study was done on colonies that had tubes removed and then were cultured in channels at four flow velocities. This experiment was designed to induce a phenotypic plastic response to flow. Instead, I found no significant difference in tube length or tube number in response to four flow velocities. This result suggests that the tube development of R. recondita may be canalized, or fixed. It is significant because it suggests that small differences that distinguish primitive, encrusting graptolite species, are good. Chapter five is on the composition of R. recondita tubes. Several hypotheses and numerous analysis have been done on this topic, but none were conclusive. Here I use genomics and bioinformatics, immunochemistry and spectroscopy and reject the hypotheses that the tubes contain keratin or cellulose. Instead I found eight chitin synthase genes in the genome and transcriptome, a complex made of a chitin-like polysaccharide, protein, fatty acid and unexpected elemental components. This study is significant because it closes the door on old hypothesis of graptolite tube composition and reveals that it is a complex structure including chitin. The conclusion chapter is a brief summary of the results and a reflection on fruitful avenues of future research.

Cette étude comparative est une révision de la famille des Harrimaniidae basée sur les caractères morphologiques d'espèces connues et nouvelles provenant des collections de William E. Ritter, Theodore H. Bullock et Kandula P. Rao rassemblées au cours du 20e siècle. Les descriptions présentées ici portent le total des genres de cinq à neuf par l'ajout de Horstia n. gen., Mesoglossus n. gen., Ritteria n. gen et Saxipendium, un genre auparavant attribué à la famille monospécifique des Saxipendidae. Le nombre d'espèces est porté à 34 par la description de cinq nouvelles espèces du Pacifique oriental: Horstia kincaidi, Mesoglossus intermedius, Mesoglossus macginitiei, Protoglossus mackiei et Ritteria ambigua. La description d'une sixième espèce, Stereobalanus willeyi Ritter et Davis, 1904 (nomen nudum) est présentée ici pour la première fois, ainsi qu'une description abrégée de Saxipendium coronatum. Quatre espèces précédemment attribuées au genre Saccoglossus sont transférées au genre Mesoglossus: M. bournei, M. caraibicus, M. gurneyi, et M. pygmaeus et Saccoglossus borealis est transféré au genre Harrimania. Une hypothèse phylogénétique sur la famille des Harrimaniidae est émise, présentant l'évolution possible des caractères morphologiques au sein du groupe. Finalement, des notes sur la distribution géographique étendue mais discontinue de plusieurs espèces suggère que les entéropneustes auraient pu avoir une distribution ancienne continue et plus grande qui aurait été fragmentée par la suite.<br>This comparative study is a revision of the family Harrimaniidae based on morphological characters of described and undescribed species from the collections of William E. Ritter, Theodore H. Bullock and Kandula P. Rao, gathered in the 20th century. The new descriptions bring the total number of genera to nine by the addition of Horstia n. gen., Mesoglossus n. gen., Ritteria n. gen and Saxipendium, a genus previously assigned to the monospecific family Saxipendidae The number of species is increased to 34, resulting from the description of five new species from the eastern Pacific: Horstia kincaidi, Mesoglossus intermedius, Mesoglossus macginitiei, Protoglossus mackiei and Ritteria ambigua. The description of a sixth species, Stereobalanus willeyi Ritter et Davis, 1904 (nomen nudum) is presented here for the first time and a brief description of Saxipendium coronatum is also presented. Four species previously assigned to the genus Saccoglossus are transfered to the genus Mesoglossus: M. bournei, M. caraibicus, M. gurneyi, and M. pygmaeus, while Saccoglossus borealis is transfered to the genus Harrimania. A phylogenetic hypothesis on the Harrimaniidae is postulated presenting the possible evolution of morphological characters within the group. Finally, notes on the wide but spotty distribution of several species suggest that the Enteropneusta may have once had a wider distribution that has since become fragmented.

博士<br>國立中興大學<br>生物科技學研究所<br>107<br>Mesoderm is a crucial germ layer that contributes to the complexity of bilaterian animals. In chordates, mesodermal cells are first specified into dorsal notochord to support the body and bilateral somites that later differentiate into muscles. These two mesodermal structures are defining features of chordates and are believed to contribute to a better motility for their tadpole-type larvae comparing to non-chordate larvae that propel by cilia. Studies in various chordate species have demonstrated that fibroblast growth factor (FGF) signaling is essential for notochord development through the activation of brachyury at notochord in vertebrates and ascidian. FGF signaling also regulates somite and muscle development in vertebrates but not the formation of trunk muscle in ascidian. In non-chordate deuterostome such as sea urchin embryos, FGF signaling is required for muscle development, but not the expression of brachyury. The functional differences of FGF signaling in controlling muscle development lead to ambiguity in the ancient role of FGF signaling in deuterostomes, and how FGF signaling had evolved to a new function in controlling notochord development through brachyury in chordates. In order to understand the role of FGF signaling during deuterostome evolution, we investigated functions of FGF signaling in mesoderm development during embryogenesis and metamorphosis in a non-chordate marine animal Ptychodera flava, an indirect-developing hemichordate that possess larval morphology similar to echinoderms and adult body features that resemble chordates. We have identified five FGF ligands and three FGF receptors in P. flava. Phylogenetic analyses revealed that hemichordates possess a conserved FGF8/17/18 in addition to several putative hemichordate-specific FGFs. Further functional studies showed that the mesodermal cell fate is specified at the early gastrula stage, and then theses cells are differentiated stepwise into the hydroporic canal, the pharyngeal muscle, and the muscle string; notably, formation of the last two muscular structures are regulated by FGF signaling. Moreover, the transcription levels of FGF ligands and receptors were significantly increased during metamorphosis, and augmentation of FGF signaling accelerated the process, suggesting its’ role in facilitating the transformation from cilia-driven swimming larvae into muscle-driven worms. These results support the ancestral role of FGF signaling in muscle development in deuterostomes. Further studies are in progress for elucidating how the novel role of FGF signaling in notochord development had evolved from its ancestral role in the lineage leading to chordates.

AbstractHemichordates are benthic marine invertebrates closely related to chordates. Several species, including Ptychodera flava in the phylum Hemichordates, can undergo whole body regeneration from a small fragment. P. flava is widely distributed in the warm Indo-Pacific region and is easily collected in the lower tidal zone of a shallow beach with a coral reef. Here, we describe the methods for animal collection and preparation of regenerating tissues. The prepared tissues can be used for various molecular and/or histological experiments. We also demonstrate how to examine gene expression patterns in the tissues using whole mount in situ hybridization.