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Characterization of the Gastro-Entero-Pancreatic System of Osteoglossomorpha: an Immunohistochemical, Immunocytochemical, and Molecular Study

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Characterization of the Gastro-Entero-Pancreatic System of Osteoglossomorpha: an Immunohistochemical, Immunocytochemical, and Molecular Study Characterization of the gastro-entero-pancreatic system of Osteoglossomorpha: an immunohistochemical, immunocytochemical, and molecular study Azza Abdul Fattah Al-Mahrouki A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy in the Department of Zoology University of toronto @ a Al-Mahrouki, 2001 National Library Bibliothèque nationale 1*1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. nie Wellington Ottawa ON KI A ON4 Ottawa ON KIA ON4 Canada Canada vow& Yom réienww Our fi& Notre réftirence The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Lïbrary of Canada to Bibliothèque nationale du Canada de reproduce, loan, distri'bute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation- Abstract Characterization of the gastro-entero-pancreatic system of Osteoglossomorpha: An immunohistochemical, immunocytochemical and molecular study. Doctor of Philosophy, 2001- Azza Abdul Fattah Al-Mahrouki. Department of Zoology. University of Toronto. The principal objectives of this study were twofold: fjrst, to compare the distrr'bution and structure of the GEP system of the osteoglossomorphs with that of the other teleosts and with vertebrates in general; second, to examine the possibility of using the primary structure of the GEP system peptides in a phylogenetic analysis. To fulnU these objectives several approaches were use& including histolo gy, immunohisto - and immunocyto chernistry, and mo lecular cloning. The results indicated that an islet organ (endocrine pancreas) with aelyscattered islets consists of four cell types (A, B, D and F); there are two D-ceU subtypes. Endocrine ce& which produce some of the same peptides as the islet cells were also observed within the intestinal epithelia; some variation occurs among the species. For exaziiple, insulin was detected in ceils wIthin the intestinal and stomach epithelia of Pantodon bucholzi, and Sequently in the intestinal epithelia of Osteoglossum biciwhosum and Gnarhonernuspetersii, but not in the other species. In the molecular biology studies, preproinsulin and the preprosomatostatin sequences were deduced fiom the corresponding cDNA sequences and were used in the phylogenetic aflSL].ysis.The phylogenetic analysis of the preproinsulin sequences confïrmed the monophyletic grouping of the osteoglossomorphs, and also indicated that the osteoglossomorphs are not the most basal living teleosts as previously thought, but rnay be more generalized teleosts. In addition, the phylogenetic analysis showed the elephantnose to be closely related to the feather fh kdefis4 and the arawana to be closely related to the butterfly fïsh. These data are consistent with previous reports (Lamier and Liem 1983; Li and Wilson 1996). The intragroup relationship was cobedby phylogenetic analysis using sequences of cDNA of preprosomatostatia These resdts indicated that the primary structure of the GEP system peptides is usefül for inferring phylogenetic analysis to some extent, but also that the sequence analysis of more than one peptide should be considered in such analyses. This study suggests that there is a generalized position of the GEP system in cornparison with that of the other teleosts. Acknowledgments No matter how independent 1 like to think myseK 1couid not have completed the Doctor of Philosophy's degree thesis without the help of mmy other people to whom 1owe at Ieast a debt of gratitude. 1would hrst like to mention my advisors, Professor John Youson, Professor David Butler, and Associate Professor David Irwin. Through the years, Prof. Youson has provided sound advice at appropriate crossroads. His enthusiasm, scient& insight, and confidence has contriibuted immensely to my research, and to the completion my thesis. Thank you for giving me the research opportunity and for the research fieedom that promoted my self confidence and autonomy. 1thank Prof. Butler for bis advice, guidance, and his very fhitful discussions on the philosophicai views of the research; it is so important to bandy ideas with more experienced researchers. 1 am also thankful to Dr. Irwin for his advice and constructive feedback in the field of molecular biology: hi, help with the editing of the last two chapters of the thesis has ken invaluable. 1 am also grateful for his help with the production of the phylogentic trees in this thesis. 1 &O want to aclmowledge Dr. E. Plketskaya, Dr. P. Andrews, and Dr. C. Yip for providing me with some of the antisera used in this research. 1 am especialiy gratefùi to Dr. L. Graham for providing me with the goldeye tissues, which otherwise have been very hard to obtain 1 also want to acknowledge Dr. M. Wilson for allowing me to use the cladograrn fkom his publication in the general introduction to this thesis. 1 would iike to thank Raymond Or for teaching me the use of the electron microscopy, his technical help is very much appreciated. The graphic expertise of Ken Jones helped in the final production of the photogmphs in the thesis, and is also very much appreciated. My coileagues at the Division of Life Sciences are all acknowledged for their advice on Metent research issues and for helping me in many different ways. Thank you to everyone, 1am extremely gratefûl. 1 also owe special thanks to my family, to whom 1 would Like to dedicate Siis thesis - to, not only for their moral support, but for bearing with me throughout this journey and creating a very positive atmosphere, making it easier to withstand the difficulties that sometimes arose. To my parents, Nabilla and Abdul Fattah Al-Mahrouki, and to my children, Ahmad and Areag Osman: thanks for helping me in every way you could. This study was supported by a grant ftom the Naturai Sciences and Engineering Research Council of Canada to Dr. John H. Youson. Abbreviations APUD amine precursor uptake and decarboxylation aPY anglerfish polypeptide tyrosine CAS casein CODEHOP Consensus-Degenerat e Hybrid Oligonucleo tide Primers GEP gastro-entero-pancreatic system GLP glucagon-like peptide GLU glucagon INS insulin mINS mammalianinsulin WY neuropeptide tyrosine 0s04 osmium tetroxide PAUP phylogenetic analysis using parsimony PBS phosphate buffered saline PCR polymerase chain reactiom PYY peptide tyrosine tyrosine RACE rapid amplification of cDN-4 ends RNA riinucleic acid SST somatostatin SST-14 synthetic invariant somato statin- 14 SST-25 sahon somatostatin-25 SST-34 iamprey somatostatin-34 Table of Contents Abstract ii Aclmowledgements iii Abbreviations v List of figures ix List of tabIes xiv General Introduction 1 Chapter 1 Tmmunohistochemical studies of the endocrine cells within the gastro-entero- pancreatic system of Osteoglossomorpha Introduction 15 Materials and Methods Anirnals Tissue preparation - Histology - Lrnmunohistocbemistry Results Morphology and histology + Immunohistochemistry - Islet cells - Gastro-intestinal cells Discussion 34 Chapter 2 hunohistochemistry of the endocrine cells in the gastro-entero-pancreatic system of a generalized teleost (Catostomus comrnersoni) and a more derived teleost (Am bloplites rupestris). Introduction Materials and Methods Animals 0 Tissue preparation - Histology - Immunohistochemistry Results 0 General morphology Immunohistochemistry - Islet ceils - Gastro-intestinal cek Discussion Chapter 3 Ultrastructure and immunocytochernistq of the islet organ of Osteoglossomorpha Introduction Materials and Methods Animais 0 Tissue preparation - Routine electron microscopy - Immynocytochemistry Results Discussion Chapter 4 Molecular cloning of preproinsulin cDNAs fiom several osteoglossomorphs and a cyprinid Introduction Materials and Methods 74 Animais a RNA extraction, first-Strand cDNA synthesis & RACE vii Isolation of the preproinsulin eDNAs Sequencing & identification of the cDNAs Results Discussion O Structure analysis O Phylogenetic anabsis Chapter 5 Characterization of somatostatin cDNAs molecular identification and comparative analysis Introduction Materials and Methods Animais Extraction and quantification of total RNA Isolation of the preprosomatostatin cDNAs & sequence anaSsis O Northern analysis 93 Results Discussion Structure heterogeneity Phylogenetic aaalysis General Discussion 106 Summary 117 Literature Cited 120 Appendix 1 1 Appendix 2 II Appendix 3 III: Glossary List of Figures Figure Page L Diagrammatic representatim of the distrr'bution of the endocrine tissue 13 in fish relative t O the exocrine pancreatic tissue. 2 Phylogenetic relationships of major osteoglossomorph taxa. 3 Diagrammatic representation of a ventral view of the viscera in fke species of osteoglossomorphs. 4 Photograph of a ventrai view of the viscera in H ahsoides. 5 Sections of pancreatic islets and exocrine acini (e) stained with hematoqdin and eosin [A] and various antkra p-Dl. 6 Sections of islets of several osteoglossomorphs demonstrating immuoreactivity to various antisera.
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