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Information to Users INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6” x 9” black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell Information Company 300 North Zeeb Road, Arm Arbor MI 4S106-1346 USA 313/761-4700 800/521-0600 CHARACTERIZATION OF A SPAWNING PHEROMONE OF PACIFIC HERRING by Joachim Schnorr von Carolsfeld B.Sc., MSc, University of Victoria A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology We accept this thesis as conforming to the required standard N.M. Sherwood, Sufiervisor (Department of Biology) G O. Mackie, Departrn^tal Member^(Department of Biology) 0. Hawryshyn, Department ber{Department of Biology) J. Ausio, Outsid artment of Biochemistry) N.E. Stacey,''Exterqgl Examiner (Department of Zoology, University of Alberta) ©Joachim Schnorr von Carolsfeld, 1997 University of Victoria All rights reserved. This thesis may not be reproduced in whole or in part, by mimeograph or other means, without the permission of the author. XI ABSTRACT Pacific herring, Clupea harengus pallasi, possesses a pheromone in the milt and testes that triggers spawning behaviour in reproductively mature individuals of both sexes, and plays a role in synchronizing the school spawning that is distinctive of this species. The pheromone was found to be effective as a transient olfactory stimulus in eliciting a behavioural response that varied in the degree of expression and time course. Stimulus strength was found to influence the time course of the response, whereas differences in maturity, evident through examination of plasma levels of steroids, were correlated with a propensity to respond to the pheromone. Input from factors other than the spawning pheromone appear to be needed to elicit prolonged spawning; some of these factors also act through olfaction. Immediate effects of stress were not found to influence the response to the spawning pheromone. Plasma levels of reproductive steroids of herring during the spawning season were measured with radioimmunoassays. Peak levels of 17,20p-dihydroxy-4-pregnen-3-one (17,20p-P) were found to coincide with final maturation in females and the initiation of milt production in males, suggesting that this steroid is the maturation-inducing steroid of this species. Other features found to be distinctive of the reproductive physiology of the herring included low plasma levels of the unconjugated maturation-inducing steroid, high levels of 17a-progesterone (17-P) and 3a,17a-dihydroxy-5p-pregnan-20-one (3a,17-P-5p), and high levels of glucuronated steroids. Structural investigation of the pheromone with liquid chromatography/ mass spectrometry showed that it consists of at least two components which do not elicit a behavioural response individually. One of these compounds is sulphated 17,20p-P. Ill The structure of proteinaceous hormones involved in controlling reproduction of the herring was also investigated. It was shown that this species possesses three forms of gonadotropin-releasing hormone (GnRH) in the brain, one with a structure that has not been reported before. These results indicate that the presence of three GnRH forms is a primitive, rather than derived, condition in the teleosts. The structure of the (3-subunit of gonadotropin II (GtH ll-P) of herring was also deduced by isolation of a cDNA for this molecule. The structure of the herring GtH ll-p was found to be quite different from other teleost molecules of this kind, and a phylogenetic analysis of known GtH ll-P structures suggests that the p-subunit of both mammalian gonadotropins may be most closely related to the p-subunit of teleost GtH-l. N.M. Sherwood, Stzpervisor (Department of Biology) G.O. Mackie, Departmental Member (Department of Biology) C. Hawryshyn, Departmental M er^pr (Department of Biology) J. Ausio. CXTtgme Menfl^ ^ B ^ ^ ftm en t of Biochemistrvl N.E. Stacey, ExternaL/fcxaminer (Department of Zoology, University of Alberta) IV TABLE OF CONTENTS ABSTRACT...........................................................................................................ii TABLE OF CONTENTS.................................................................................... iv LIST OF TABLES............................................................................................. viii LIST OF FIGURES............................................................................................. ix LIST OF ABBREVIATIONS........................................................................... xiiii ACKNOWLEDGEMENTS.................................................................................xv CHAPTER 1: OVERVIEW OF REPRODUCTIVE PHEROMONES AND THE ROLE OF HERRING RESEARCH................................................ 1 REFERENCES CITED.................................................................................................13 CHAPTER 2. BEHAVIORAL CHARACTERIZATION OF PHEROMONE-INDUCED SPAWNING OF PACIFIC HERRING 24 ABSTRACT...................................................................................................................24 INTRODUCTION..........................................................................................................25 MATERIALS AND METHODS...................................................................................27 SPECIFIC METHODS AND RESULTS................................................................... 29 A) CHARACTERIZATION OF THE PHEROMONE RESPONSE:........... 29 8) FACTORS INFLUENCING THE RESPONSE TO THE SPAWNING PHEROMONE:................................................................44 V DISCUSSION................................................................................................................ 64 REFERENCES CITED................................................................................................ 72 CHAPTER 3. REPRODUCTIVE STEROIDS DURING MATURATION IN THE PACIFIC HERRING (CLUPEA HARENGUS PALLASI).................................................................................... 76 ABSTRACT....................................................................................................................76 INTRODUCTION...........................................................................................................77 MATERIALS AND METHODS....................................................................................78 RESULTS...................................................................................................................... 86 DISCUSSION.............................................................................................................. 109 REFERENCES CITED.............................................................................................. 125 CHAPTER 4. PLASMA STEROIDS DISTINCTIVE TO HERRING RESPONSIVE TO THE SPAWNING PHEROMONE.............................. 135 ABSTRACT..................................................................................................................135 MATERIALS AND METHODS..................................................................................137 RESULTS.................................................................................................................... 142 DISCUSSION.............................................................................................................. 143 REFERENCES CITED.............................................................................................. 152 CHAPTER 5. IDENTIFICATION OF A COMPONENT OF THE SPAWNING PHEROMONE OF THE PACIFIC HERRING BY LIQUID CHROMATOGRAPHY / MASS SPECTROMETRY (LC/MS).............................................................................................................. 155 VI ABSTRACT..................................................................................................................155 INTRODUCTION........................................................................................................ 156 MATERIALS AND METHODS..................................................................................159 RESULTS....................................................................................................................184 DISCUSSION..............................................................................................................205
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