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The Original Manuscript Received by UMI Contains Pages with Indistinct Print. Pages Were Microfilmed As Received. This Reproduction Is the Best Copy Available NOTE TO USERS The original manuscript received by UMI contains pages with indistinct print. Pages were microfilmed as received. This reproduction is the best copy available UMI EVOLUTION OF THREE NEUROPEPTIDES ISOLATED FROM THE BRAIN OF STURGEON David William Lescheid B.Sc., Simon Fraser University, 1989 A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology We accept this dissertation as conforming to the required standard rwood (Dept, of Biology) Dr.D. Burke (Dept of Biology) Dr. S. Misra (Dept of Biochemistry and Microbiology) Dr.'T. W. Pearson (Dept, of Biochemistry and Microbiology) Dr. E. M. Donaldson (External examiner) © David William Lescheid, 1997 University of Victoria All rights reserved. This dissertation may not be reproduced in whole or in part, by photocopying or other means, without permission from the author. 11 ABSTRACT In vertebrates the brain superimposes control on fundamental processes such as reproduction and growth. Neuropeptides secreted from the brain initiate a cascade of events that affect these processes. In this thesis three neuropeptides are examined to determine their structures and patterns in the context of vertebrate evolution. Reproduction in vertebrates is controlled by the neuropeptide gonadotropin- releasing hormone, GnRH, a decapeptide belonging to a peptide family of twelve known members. One common theme in vertebrates is that there is usually more than one form of GnRH in the brain of a single species; often each form of GnRH has a separate location in the brain and therefore, an implied distinct function. In this thesis, the brain of Siberian sturgeon, Acipenser gueldenstaedti, initially was examined for GnRH using reversed- phase high performance liquid chromatography, HPLC, and radioimmunoassay, RIA, with specific antisera and was shown to contain mammalian (m)GnRH by chemical sequence analysis and by accurate determination of the molecular mass. In addition, another form of GnRH, termed chicken (c)GnRH-ll, was found in the sturgeon brain. This is the first report to show that the primary structure of GnRH is identical in an evolutionarily-ancient fish and in mammals including humans. Further, the second form of GnRH, cGnRH-11, was identified for the first time in the brain of adult stumptail monkeys {Macaca speciosa) as well as in adult and fetal rhesus monkey {Macaca mulatta) brains. This study implies that at least two forms of GnRH are found in the brain of most vertebrate species including mammals. In cartilaginous fish that evolved earlier than sturgeon, the same HPLC and RIA methods were used to demonstrate that regions of the brain and pituitary of skate. Raja canebensis, also contained cGnRH-11 but dogfish (df)GnRH rather than mGnRH. By the same criteria, teleost fish like whitefish {Prosopium williamsoni), platyfish {Xiphophorus maculatus), green swordtail {Xiphophorus hellerei) and sablefish {Anoplomia fimbria) Ill were shown to have cGnRH-II and salmon (s)GnRH, as well as one or two more immunoreactive variants of GnRH with novel or seabream (sb)GnRH-like properties, within their brain. The identity of at least three types of immunoreactive GnRH molecules in the brain of these fish species suggests that three forms of GnRH in the brain is an early condition in teleost evolution. Ancestral sturgeon emerged at a branch point between the bony fish lineage and the tetrapod lineage and therefore, it is useful to compiate the neuropeptide structures found in their brain with those both in fish and more evolutionarily-advanced vertebrates. Several tetrapod species were examined to determine if the forms of GnRH found in the sturgeon brain had been retained in their evolution. In contrasts to studies in our laboratory and by others showing that most amphibians, reptiles and birds contain two forms of GnRH, the present research shows that the brain of the green anole lizard, Anolis carolinensis, contained only cGnRH-11 within its brain. In addition, my HPLC and RIA studies showed that only mGnRH was present in the brain of guinea pig, hamster and rat suggesting that there are some species which function with only one form of GnRH in their brain. Also, there were no distinguishable forms of GnRH in a human placenta, demonstrating that the type(s) of GnRH might be tissue-specific. Two neuropeptides associated with growth also were isolated from the sturgeon brain. A cDNA encoding growth hormone-releasing factor, GRF, and pituitary adenylate cyclase-activating polypeptide, PACAP, was isolated and sequenced using the polymerase chain reaction, PGR, and other molecular biology methods. In contrast to mammals where GRF and PACAP are encoded on separate genes, in sturgeon, GRF and PACAP are encoded in tandem on a single mRNA. IV In this thesis, I establish the structure of GnRH, GRF, and PACAP in sturgeon, a species that evolved near a critical branching point between bony fish and tetrapods. These structures are used as a focal point for comparison to those of other vertebrates. This comparative evolutionary approach is an important step toward understanding the evolution of these important neuropeptides as well as enhancing our knowledge of general principles in the endocrine systems controlling reproduction and growth. Examiners: Dr. N (^ . ood (Dept, of Biology) Dr. R. D. Burke (Dept, of Biology) Dr. S^^Misra (Dept of Biochemistry and Microbiology) D r.' '. W. Pearson (Dept, of Biochemistry and Microbiology) Dr. E.M. Donaldson (External examiner) TABLE OF CONTENTS ABSTRACT......................................................................................................ii TABLE OF CONTENTS................................................................................... v LIST OF FIGURES ........................................................................................viii LIST OF TABLES............................................................................................ xii LIST OF ABBREVIATIONS.......................................................................xiv ACKNOWLEDGEMENTS........................................................................... xvi CH A PTER I: General Introduction ..................................................................1 GnRH peptide structure ............................................................................8 GnRH; function and location .................................................................. 11 GnRH mRNA and gene structure ...........................................................12 GRF peptide structure ..............................................................................16 GRF: location and function ..................................................................... 19 GRF mRNA and gene structure ..............................................................19 PACAP peptide stru ctu re .........................................................................21 PACAP: location and function ..................................................................21 GRF-like/ PACAP and PACAP mRNA and gene structure ..................23 Glucagon superfamily .............................................................................. 24 Transport of GnRH, GRF and PACAP in the brain of vertebrates 25 GnRH receptor .......................................................................................... 26 GRF and PACAP receptor superfamily ................................................. 27 GRF receptor ............................................................................................. 27 PACAP receptor ........................................................................................28 Overlapping functions ...............................................................................29 Prevertebrate history ...................................................................................30 Purpose of thesis ........................................................................................ 30 Literature cited ............................................................................................ 33 VI CHAPTER 2: Determination of the primar\' structure for mGnRH from the brain of Siberian sturgeon, Acipenser gueldenstadtii ......................47 Introduction ...............................................................................................48 Materials and M ethods ............................................................................... 50 R esults......................................................................................................... 55 D iscussion ...................................................................................................65 Literature c ite d ..............................................................................................68 CHAPTER 3: Identification of the forms of GnRH present in the brain of a cartilaginous fish, the skate, as well as in the brains of four teleosts: whitefish, platyfish, green swordtail and sablefish ..............................73 Introduction ................................................................................................ 74 Materials and M ethods .................................................................................78 R esults...........................................................................................................88 D iscussion ................................................................................................... 112 Literature
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