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Canada Du Canada Nallonal Ubrary 81blloth~"Que n3tlon31e of Canada du Canada Acquisitions and Direction des acq J1Sliions cl 81bllOgrç;phlC ServIces Branch des services blbllU\]raph\Qucs 395 Welllnqton StrL'c! 3~5. rUt,' \\'clhnqton OnJw,J, On:;1rIO OI:.lW~l (On13f10) K1AON': K1AON·; " ~ " ',.,' ,",' " " " .'.., , " "'.' " ''''''''', " NOTICE AVIS The quality of this microform is La qualité de cette microforme heavily dependent upon the dépend grandement de la qualité quality of the original thesis de la thèse soumise au submitted for microfilming. microfilmage. Nous avons tout Every effort has been made to fait pour assurer une qualité ensure the highest quality of supérieure de reproduction. reproduction possible. If pages are missing, contact the S'il manque des pages, veuillez university which granted the communiquer avec l'université degree. qui a conféré le grade. Sorne pages may have indistinct La qualité d'impression de print especially if the original certaines pages peut laisser à pages were typed with a poor désirer, surtout si les pages typewriter ribbon or if the originales ont été university sent us an inferior dactylographiées à l'aide d'un photocopy. ruban usé ou si l'université nous a fait parvenir une photocopie de quamé; inférieure. Reproduction in full or in part of La reproduction, même partielle, this microform is governed by de cette microforme est soumise the Canadian Copyright Act, à la Loi canadienne sur le droit R.S.C. 1970, c. C-30, and d'auteur, SRC 1970, c. C-30, et subsequent amendments. ses a.nendements subséquents. Canada 1 • Characterization of a protein binding site involved in the regulation of transcription elongation within the murine c--mye gene by DANIEL DUFORT Department of Medicine Division of Experimental Medicine McGill University Montreal, Quebec December 1993 A Thesis submitted to the Faculty of Graduate Studies and Research in partial fulfilment of the requirements of the degree of Doctor of Philosophy © Daniel Dufort 1993 • National Ubrary 8lbltoth~que nationale 01 Canada du Canada AcquIsitions and DIrection des acqwslllons ct Bibliographie Services Branch des services bibliographiques 395 Weill; IQlon Streel 395. rue Wellington Ottawo..Ontano On;lwa (Ont~Hlo) K1A ON4 KlA ON·1 The author has grc:nted an L'auteur a accordé une licence irrevocable non-exclusive licence irrévocable et non exclusive allowing the National Library of permettant à la Bibliothèque Canada to reproduce, loan, nationale du Canada de distribute or sen copies of reproduire, prêter, distribuer ou his/her thesis by any means and vendre àes copies de sa thèse in any form or format, making de quelque manière et sous this thesis available to interested quelque forme que ce soit pour persons. mettre des exemplaires de cette thèse à la disposition des personnes intéressées. The author retains ownership of L'auteur conserve la propriété du the copyright in his/her thesis. droit d'auteur qui protège sa Neither the thesis nor substantial thèse. Ni la thèse ni des extraits extracts from it may be printed or substantiels de celle-ci ne otherwise reproduced without doivent être imprimés ou his/her permission. autrement reproduits sans son autorisation. ISBN 0-3IS-94613-X Canada Ac-mye promoter element involved in transcription elongation Il • Abstract The c-mye gene was previously shown to be regulated by a condilional block to transcription elongation and sequences from its promoter were implicated in this regulation. The objective of this project was to define the promoter elements involved in the control of transcription elongation. Using heterologous promoter constructs, the MElal protein binding site located in the c-mye promoter was shown to be required for the block to transcription elongation. From mutation analysis, a correlation was established between the binding of nuclear factors to MElal sites in vitro and the ability of these sites to confer block to transcription elongation in vivo, strongly implicating trans-acting factors in this process. Fractionation studies demonstrated that three nudear factors interact with the MElal site, thus generating three protein-DNA complexes termed "a", "b", and "c". These factors were characterized and a cDNA encoding the protein responsible for complex "c" was isolated. This protein was shown to represent the human homologue of the Drosophila Cut homeodomain protein (hu-Cut) and to repress expression from the c-mye promoter in transient transfection assays. • 111 • Résumé L'expression du gène c-mye est régulée en partie au niveau de l'élongation de la transcription. Le but de ce projet était de définir les éléments du promoteur impliqués dans le contrôle de l'élongation de la transcription. J'ai montré que le blocage de la transcription était augmenté lorsque le site MElal, une séquence de 25 pb présente dans le promoteur de c­ mye, était introduit dans des constructions contenant un promoteur hétérologue lié à l'exon 1 de c-mye. Une analyse mutationnelle a permis d'établir une corrélation entre l'interaction avec des facteurs nucléaires in vitro et la capacité de conférer le blocage de la transcription in vivo. Ces résultats suggéraient que des facteurs de transcription se liant au site MElal pouvaient jouer un rôle dans le contrôle de l'élongation. Des études de fractionnement ont révélé que trois facteurs nucléaires pouvaient interagir avec le site MElal pour former les complexes "a", ''b'' et "c", Ces facteurs ont été analysés et le cDNA codant pour la protéine formant le complexe "c" a été isolé. J'ai montré que cette protéine était i'homologue humain de l'homéoprotéine Cut de la drosophile et qu'elle pouvait réprimer l'expression de gènes rapporteurs contenant le site MElal ainsi que l'exon 1 de c-mye. • 1 \' • Acknowledgments 1 would like to thank my wife Celine and my little girl Alexandra for their love, understanding, and patience throughout these studies. 1 wish to express my gratitude to my supervisor, Dr, Alain Nepveu, for his advise and for giving me the opportunity to do this work. 1 would aIse like to thank the members of the Nepveu lab for their stimulating discussions and advise as weIl as for their friendship. Finally, 1 wish to thank the National Cancer Institute of Canada for financial support during these studies. • v • Preface The Guidelines Concerning Thesis Preparation Issued By The Faculty Of Graduate Studies And Research At McGill University reads as follows: ''The candidate has the option, subject to the approval of their department, of including as part of the thesis, copies of the text of a papers(s) submitted for publication, or clearly-duplicated text of a published paper(s), provided that these copies are bound as an integral part of the thesis. If this option is chosen, connecting texts, providing logical bridges between different pages, are mandatory. The thesis must conform to all other requirements of the "Guidelines Concerning Thesis Preparation" and should be in a literary form that is more than a mere collection of manuscripts published or to be published. The thesis must include, as separate chapters or sections: (1) a Table of Contents, (2) a general abstract in English and French, (3) an introduction which clearly states the rationale and objectives of the study, (4) comprehensive general review of the background literature to the subject of the thesis, when this review is appropriate, and (5) a final conclusion and/or summary. Additional material (procedural and design data, as weIl as descriptions of equipment used) must be provided where appropriate and in sufficient detail (e.g.. in the appendices) to allow a clear and precise judgment to he made of the importance and originality of the research • reported in the thesis. YI In the case of manuscripts co-authored by the candidate and others, the • candidate is required to make an explicit statement in the thesis of who contributed to such work and to what extent; supervisors must attest to the accuracy of such daims at the Ph.D. Oral Defense. Since the task of the Examiners is made more difficult in these cases, it is in the candidate's interest to make perfectly dear the responsibilities of the different authors of co-authored papers." l have chosen to write my thesis according to to the above quoted option with one paper published and two submitted for publication. The thesis is organized into five chapters. Chapter l is a general introduction and literature review with references. Chapters II-IV contain furee manuscripts, each with its own abstract, introduction, methods, results and references. Chapter V is a general discussion of ail the results with references. Severa! people have contributed to the work presented in this thesis. In the second cb.apter, "A Protein Binding Site from the Mutine C-Myc Promoter Contributes to Transcriptional Block", Marc Drolet made the MElal-Sac construct and we both obtained stably transfected ceU population and we both performed the nueler run-on assays. Also, the MEC-MElal+ and MEC-MElal­ constructs were made by Harvey Miller, and the cell populations were established by Alain Nepveu who performed the nuelear run-on assays as weil as the SI mapping of the cytoplasmic RNA for these constructs. In the third chapter, ''The human Cut homeodomain protein represses transcription fron the c-myc promoter~', the MElal-eAT construct was made by Ryoko Harada, and the MElal-El-eAT construct was made by Ginette • Bérubé. Vil • Contributions to original research 1. The MElal protein binding site located in the promoter of the c-mye gene was shown to be important for transcriptional block. 2. A correlation was established between the ability of the MElal site to bind cellular factors in vitro and the ability to confer block to transcription elongation in vivo, strongly suggesting that cellular factors are involved in transcriptional block.
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