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University of Alberta University of Alberta GENE EXPRESSION IN BLASTOCYSTS AND THE ENDOMETRIUM DURING PORCINE PERI-IMPLANTATION DEVELOPMENT Linghuo Jiang O A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy Animal Science Department of Agricultural, Food and Nutritional Science Edmonton, Alberta Fall, 1999 National tibrary Bibliothèque nationale 1+1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. rue Wellington Ottawa ON K 1A ON4 OnawaON KlAONQ Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive Licence aliowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur format électronique. The author retains ownershp of the L'auteur conserve la propriété du copyrÏ&t in this 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 othenvise de celle-ci ne doivent être imprUnés reproduced without the author's ou autrement reproduits sans son permission. autorisation. my de,my daughter, my parents, and my parents-in-law for their constant support during the course of this study. ABSTRACT Macrophage colony-stimulating factor (CSF-1) was found to be the major factor, in conditioned media of Jag-1 cells, responsible for stimulating porcine macrophage proliferation in vitro. A porcine CSF-1 cDNA was isolated and characterized from Jag-1 cells. The CSF-1 gene was found to be present as a single copy in the porcine genome. Expression of CSF-1 transcripts was up regulated in both blastocyst and endometriai tissues during the peri-implantation period in the pig, with higher levels in endometrial tissues than blastocyst tissues. Similarly, tiigher levels of c-fms (CSF-1 receptor) transcripts were detected in endometrial tissues than in blastocysts. Our data suggest that CSF-1 might exert both paracrine and autocrine effects on porcine pre- implantation embryonic development. Expression of matrix metalloproteinase (MMP) transcripts and their regulators was examined in pregnant gilts dunng the peri-implantation penod. In blastocysts, MMP- 2, MMP- 14 and TIMP-2 increased, wfiile MMP-9 was undetectable. MMP-8, TNP-1 and TIMP-3 transcripts also increased, whereas uPA exhibited a biphasic pattern of expression. In the endometrium, expression of MMP-1, MMP-2, MMP-8, MMP-9, MMP- 1 1, MMP- 14, uPA, TIMP- 1, Tm-2 and TIMP-3 were al1 observed, but not MMP-7 and MMP-13. ProMMP-2 and Tm-2, but not proMMP-9, were detected in uterine flushings. A lack of MMP-9 expression in blastocysts might explain the non- invasive implantation seen in pigs. Co-expression of MMP-2, TIMP-2 and MMP-14 in blastocysts might contribute to extraembryonic mesoderm ceil migration. Expression of integrin subunit (al, a2,a3, a4. a5, av, pl and P3) transcripts was also determined during early gestation in pigs. In endometrial tissues, constant expression of av, Pl and P3 was detected, whereas al, a2, a3 and a5 appeared to be developmentally regulated. Expression of a2, a3, av, pl and 83 exhibited similar patterns and was developmentally regulated in blastocysts, with high levels in 1-2 mm spherical blastocysts, low levels in 5-10 mm spherical blastocysts and variable levels in elongated blastocysts. In Day-28 placental tissue, moderate expression of a 1, aS, av and P3 and weak expression of a2 and B 1 was observed. Jag-1 celIs expressed high levels of al, a2, a3, av and 83 and low levels of a5 and PL. Our findings support an important role for integrins in the processes of early ernbryogenesis, embryo implantation and placentation in the pig. in surnmary, the present study has identified and characterized the expression of some important protein factors, in blastocysts and the endometrium, that are hypothesized to play critical roles in conceptus development during early gestation in the pig. ACKNOWLEDGEMENTS First 1 would like to thank Dr. W.T. Dixon for giving me this opportunity to complete my Ph.D. study in his lab and fulfill my dream in the area of research 1 have been interested in. 1 am especially grateful for his excellent supervision as well as his emotional support and encouragement at the worst of times during the course of this sîudy. Because of bis kindness and wisdom, my life as a Ph.D. student has turned to be an enjoyable leaming experience. Special thanks are aiso given to Dr. G. R. Foxcroft for his guidance and excellent instruction in the area of reproductive biology. My sincere appreciation also goes to Dr. L.J. Guilbert for his supportive service on my Supervisory Cornmittee. Furthemore, I would like to thank Renate Meuser and Joan Turchinsky for their generous technical support al1 the way through this study. Special thanks go to Susan Novak for her help in collection of uterine and embryonic tissue sarnples, and to Dr. Denis Balcerzak, Richard Y.Z. Chai, Dr. Brian Treacy and Colin Strauss for their valuable advice and stimulating discussion. in addition, 1 want to thank ail members of Dr. G.R. Foxcroft's research group and al1 members at the MoIecular Biology and Biotechnology center for their friendship and the fun times we had in the laboratory. Finally, this study would not be possible without the support of the 1995/1996 VP Academic (Ms. Monika Lozinska) of the GSA, the previous departmental Graduate Comrnittee Chair (Dr. John Feddes), and two of my previous supervisory committee members (Drs. Kenneth Roy and Andrew Keddie). Of course, 1 should also thank Dr. Chuji Hiruki for his generous two-and-half-year financial support from NSERC for my technical and ESL training in his laboratory. TABLE OF CONTENTS LITERATURE REVIEW ......................................................................................................... .................. 1 INTRODUCTION ................................................................................................................................ 1 EARLY EMBRYOMC DEVELOPMENT iN Ti-E PIG .................................................................... 2 A . Cleavage .......................................................................................................................... 2 B . Compaction and Blastocyst Formation ............................................................................ 3 C . Formation of Germ Layers and Extraembryonic Membranes .......................................... 5 MATERNAL RECOGNITION OF PREGNANCY ........................................................................... 7 BLASTOCYST ELONGATION AND MPLANTATION IN THE PIG ................... ... ................ 9 A . BIastocyst Expansion and Elongation ............................................................................. 9 B . Blastocyst Attachment/'implantation ............................................................................... 10 POSSIBLE MECHANISMS REGULATING BLASTOCYST ELONGATION AND iMPLANTATiON LN THE PIG ......................................................................................................... 12 A . Steroids and Their Receptors .......................................................................................... 13 B . RoIes of Retinol and Retinoic Acid in Embryonic Morphorgenesis ............................... 15 C. Trophoblast-Uterine Epithelium Adhesion ................................................................ 16 D . Fonnation of Porcine Non-Invasive Placenta .................................................................20 E . Regulators of Implantation: Growth Factors and Cytokines ........................................ -24 I . Insulin-like growth factor family ....................................................................... 25 2 . Epidennal growrh factor family ........................................................................26 3. Transforming growth factor flfamily ............................................................... -28 4 . Inte$eron family ................................................................................................ 30 5. Macrophage colony-stimularing factor ............................................................. 31 6. Leukemia inhibitory factor ................................................................................ 32 7. Inrerleukin- l family ........................................................................................... 34 F. Other Factors Secreted in the Utems during Pen-implantation Period ............................ 36 f . Urerofem'n.......................................................................................................... 36 2 . Kallikrein, kininogen adkinin sysrem ............................... ,.. ............................. 36 POSSIBLE SIGNAL TRANSDUCTION PATHWAYS INVOLVED IN IMPLANTATION ......... 37 A . Signal Transduction via Intracellular Receptors ............................................................. 37 B . Signal Transduction via Cell Surface Receptors ................... .. ...................................... 39 I . Ras-MAPK parhway ...................................................................................... 39 2 . TheJAK-STATparhway .................................................................................
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