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ASIEH RASOOLIZADEH TRANSCRIPTIONAL ANALYSIS OF TRANOSEMA ROSTRALE ICHNOVIRUS (TrIV) GENES, WITH EMPHASIS ON THE REP GENE FAMILY Mémoire présenté à la Faculté des études supérieures de l’Université Laval dans le cadre du programme de maîtrise en biologie pour l’obtention du grade de Maître ès Sciences (M.Sc.) DÉPARTEMENT DE BIOLOGIE FACULTÉ DES SCIENCES ET DE GÉNIE UNIVERSITÉ LAVAL QUÉBEC 2009 © Asieh Rasoolizadeh, 2009 RÉSUMÉ La guêpe endoparasitoïde Tranosema rostrale transmet un ichnovirus (“TrIV”) à son hôte lépidoptère, Choristoneura fumiferana, au moment de la ponte. Ce virus, lequel possède un génome segmenté d’ADNdb et ne peut se répliquer que dans l’ovaire du parasitoïde, est essentiel à la survie de la guêpe immature à l’intérieur de son hôte. Dans une étude antérieure, 86 cadres de lecture ouverts (ORF) ont été identifiés dans le génome de TrIV, dont 35 qui ont pu être affectés à des familles de gènes ichnoviraux connues. La balance n’affichait aucune similitude à des gènes connus. Dans le but d’évaluer (i) la précision de l'annotation du génome de TrIV et (ii) l'importance relative de chaque famille de gènes dans le succès du parasitisme par T. rostrale, une analyse transcriptionnelle de type qPCR a été réalisée chez des larves de C. fumiferana infectées ainsi que dans des ovaires de T. rostrale. Alors que la majorité (91%) des ORF attribués à des familles de gènes connues ont produit des transcrits dans les larves infectées, mais à des niveaux très variables, cette proportion était plus faible (67%) pour un échantillon de 12 ORF non-attribués. Parmi les sept familles de gènes présentes dans le génome de TrIV, la famille rep est la mieux représentée, avec 17 membres; tous se sont avérés être exprimés dans des larves infectées et/ou les ovaires de guêpe. Dans les chenilles infectées, cependant, les transcrits de deux d'entre eux, F1-1 et F1-2, étaient beaucoup plus abondants que ceux des autres gènes rep. De plus, le profil transcriptionnel de la famille rep était clairement différent dans les ovaires de guêpe, où le gène C166-1 a génére le plus abondant des transcrits rep, ce qui suggère que différents membres de cette famille pourraient avoir des fonctions spécifiques dans chaque hôte. L'abondance relative des segments génomiques était plus élevée pour les deux segments portant les trois gènes rep les plus fortement exprimés chez des chenilles infectées, mais la corrélation entre ces deux variables était faible pour les autres gènes rep, suggérant que des facteurs additionnels sont impliqués dans la régulation de l'expression des gènes rep chez les larves infectées. Des différences entre les gènes rep de TrIV ont également été observées en ce qui a trait à l'abondance relative des transcripts dans différents tissus de C. fumiferana, ce qui suggère l’existence de rôles distincts ou d’une spécialisation pour chacun des membres de cette famille à l’intérieur de différents tissus. Lorsqu’on compare les niveaux de transcripts rep, dans des chenilles infectées, à ceux de gènes appartenant à d'autres familles connues du génome de TrIV, un gène de la famille TrV (TrV1) et un gène rep (F1-1) se sont avérés beaucoup plus fortement transcrits que tous les autres gènes examinés, soulignant l'importance probable de ces deux familles dans la subjugation de C. fumiferana par T. rostrale. Dans les ovaires de guêpe, le profil transcriptionnel était dominé par un gène rep et par un membre d'une famille nouvellement décrite et identifiée parmi des ORF qui n’avaient pu être attribués à des familles connues; ces gènes codent pour les protéines sécrétées affichant un nouveau motif cystéine. ABSTRACT The endoparasitic wasp Tranosema rostrale transmits an ichnovirus (“TrIV”) to its lepidopteran host, Choristoneura fumiferana, during parasitization. This virus, which has a segmented dsDNA genome and can replicate only in the wasp’s ovaries, is essential to the survival of the immature wasp within its host. In a prior study, 86 putative open reading frames (ORFs) were identified in the TrIV genome, including 35 that could be assigned to previously recognized ichnoviral gene families. The balance displayed no similarity to known genes. In an effort to assess (i) the accuracy of the TrIV genome annotation and (ii) the relative importance of each gene family in the success of parasitism by T. rostrale, a temporal and tissue-specific qPCR transcriptional analysis was conducted in infected C. fumiferana hosts and T. rostrale wasp ovaries. The majority (91%) of putative ORFs assigned to known gene families were observed to be expressed in infected larvae, albeit at widely varying levels, but this proportion was lower (67%) for a sample of 12 unassigned ORFs. Among the seven known gene families present in the TrIV genome, the rep family is the numerically most important one, with 17 members; all of these were shown to be expressed in infected larvae and/or wasp ovaries. In infected caterpillars, however, two of them, F1-1 and F1-2, had much more abundant transcripts than the others. The rep transcriptional profile was markedly different in wasp ovaries, where the C166-1 gene generated the most abundant rep transcripts, suggesting that different members of this family may have host-specific functions. Relative abundance of genome segments was highest for the two segments bearing the three most highly iii expressed rep genes, but the correlation between these two variables was poor for the other rep genes, suggesting that some other factors are involved in the regulation of rep gene expression in infected larvae. Inter-gene differences were also observed in the relative abundance of TrIV rep transcripts in different C. fumiferana tissues, pointing to tissue-specific roles or specialized functions for individual members of this gene family. In comparing rep transcript levels to those of genes belonging to other known TrIV gene families, a TrV (TrV1) and a rep (F1-1) gene clearly outnumbered all other genes examined in infected caterpillars, pointing to the likely importance of these two gene families in host subjugation by T. rostrale. In wasp ovaries, the transcriptional profile was dominated by a rep gene and a member of a newly described family identified among previously unassigned ORFs; these genes encode secreted proteins displaying a novel cysteine motif. iv AVANT-PROPOS – FOREWORD During my graduate studies, I have met several people at Laval University who have shared their knowledge and experience with me to make my work both possible and more pleasant. I take this opportunity to thank them all from the bottom of my heart. More specifically, I would like to thank my co-supervisor, Dr. Michel Cusson. He generously welcomed me to his lab, gave me the opportunity to develop my competences, and provided me with scientific training on a daily basis. Dr. Cusson has a distinct way of dealing with problems and gives his students an opportunity to discover themselves and recognize their abilities. It has always been a great pleasure to share with him new results, and his constant cheering, interest and enthusiasm allowed me to push through and get through several difficult tasks. I would also like to thank my director, Prof. Conrad Cloutier, for taking time to assess my manuscript and help me get through the Master’s program. I will never forget the first course I took with him, which he (naturally) gave in French, a lovely language that, unfortunately, I do not fully grasp yet; he patiently helped me throughout the semester. Furthermore, I would like to express my gratitude to the members of our laboratory at the Laurentian Forestry Centre (LFC). In particular, I thank Catherine Béliveau and Don Stewart, two molecular biologists who have helped me by providing valuable and friendly guidance during my stay at LFC. I will also be eternally grateful for the support I received from the few real friends I made at Laval University; their friendly support was much appreciated, and I sincerely thank them all. Last, but not least, I wish to express my profound gratitude to my parents. Although I am living far away from them, they are always in my heart. The distance did not keep them from providing invaluable advice and generous support. I have always benefited from their gracious words and encouragements, which allowed me, during hard times, to keep moving forward and continue on my career path. v TABLE OF CONTENTS RÉSUMÉ…………………………………………………………………………………………ii ABSTRACT………….. ............................................................................................................... iii AVANT-PROPOS – FOREWORD .............................................................................................. v TABLE OF CONTENTS ............................................................................................................. vi LIST OF FIGURES...................................................................................................................... ix LIST OF TABLES ....................................................................................................................... xi CHAPITRE 1….INTRODUCTION ........................................................................................... 1 1.1 La tordeuse des bourgeons de l’épinette, Choristoneura fumiferana ............................ 2 1.1.1 Cycle Vital................................................................................................................. 2 1.2 Les parasitoïdes.............................................................................................................
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  • Ichneumonidae (Hymenoptera) As Biological Control Agents of Pests

    Ichneumonidae (Hymenoptera) As Biological Control Agents of Pests

    Ichneumonidae (Hymenoptera) As Biological Control Agents Of Pests A Bibliography Hassan Ghahari Department of Entomology, Islamic Azad University, Science & Research Campus, P. O. Box 14515/775, Tehran – Iran; [email protected] Preface The Ichneumonidae is one of the most species rich families of all organisms with an estimated 60000 species in the world (Townes, 1969). Even so, many authorities regard this figure as an underestimate! (Gauld, 1991). An estimated 12100 species of Ichneumonidae occur in the Afrotropical region (Africa south of the Sahara and including Madagascar) (Townes & Townes, 1973), of which only 1927 have been described (Yu, 1998). This means that roughly 16% of the afrotropical ichneumonids are known to science! These species comprise 338 genera. The family Ichneumonidae is currently split into 37 subfamilies (including, Acaenitinae; Adelognathinae; Agriotypinae; Alomyinae; Anomaloninae; Banchinae; Brachycyrtinae; Campopleginae; Collyrinae; Cremastinae; Cryptinae; Ctenopelmatinae; 1 Diplazontinae; Eucerotinae; Ichneumoninae; Labeninae; Lycorininae; Mesochorinae; Metopiinae; Microleptinae; Neorhacodinae; Ophioninae; Orthopelmatinae; Orthocentrinae; Oxytorinae; Paxylomatinae; Phrudinae; Phygadeuontinae; Pimplinae; Rhyssinae; Stilbopinae; Tersilochinae; Tryphoninae; Xoridinae) (Yu, 1998). The Ichneumonidae, along with other groups of parasitic Hymenoptera, are supposedly no more species rich in the tropics than in the Northern Hemisphere temperate regions (Owen & Owen, 1974; Janzen, 1981; Janzen & Pond, 1975), although