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Embryonic Resorption – a Longitudinal Ultrasonographic Study in the Model Species European Brown Hare (Lepus Europaeus PALLAS, 1778)

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Embryonic Resorption – a Longitudinal Ultrasonographic Study in the Model Species European Brown Hare (Lepus Europaeus PALLAS, 1778) Aus dem Leibniz-Institut für Zoo- und Wildtierforschung (IZW) im Forschungsverbund Berlin e.V. und dem Fachbereich Veterinärmedizin der Freien Universität Berlin Embryonic resorption – a longitudinal ultrasonographic study in the model species European brown hare (Lepus europaeus PALLAS, 1778) Inaugural-Dissertation zur Erlangung des Grades eines Doktors der Veterinärmedizin an der Freien Universität Berlin vorgelegt von Katharina Dorothea Ilse Lidy Schröder Tierärztin aus Berlin Berlin 2014 Journal-Nr.: 3696 Gedruckt mit Genehmigung des Fachbereichs Veterinärmedizin der Freien Universität Berlin Dekan: Univ.-Prof. Dr. Jürgen Zentek Erster Gutachter: Univ.-Prof. Dr. Heribert Hofer Zweiter Gutachter: Prof. Dr. Sabine Meinecke-Tillmann Dritter Gutachter: PD Dr. Kerstin Müller Deskriptoren (nach CAB-Thesaurus): Amniocentesis, amnion, amniotic fluid, biopsy, blastocyst, conceptus, corpus luteum, chorion, embryology, embryo mortality, embryonic development, embryonic resorption, fertility, fetal development, fetal death, fetal growth, fetal membranes, fetal resorption, hares, Lagomorpha, Lepus europaeus, Leporidae, luteolysis, mammals, ovaries, ovulation, pregnancy, pregnancy diagnosis, prenatal development, prenatal diagnosis, rabbits, regression, reproduction, resorption, trophoblast, ultrasonography, ultrasound, chorionic villi (MesH) Tag der Promotion: 15.09.2014 Bibliografische Information der Deutschen Nationalbibliothek Die Deutsche Nationalbibliothek verzeichnet diese Publikation in der Deutschen Nationalbibliografie; detaillierte bibliografische Daten sind im Internet über <http://dnb.ddb.de> abrufbar. ISBN: 978-3-86387-543-5 Zugl.: Berlin, Freie Univ., Diss., 2014 Dissertation, Freie Universität Berlin D 188 Dieses Werk ist urheberrechtlich geschützt. Alle Rechte, auch die der Übersetzung, des Nachdruckes und der Vervielfältigung des Buches, oder Teilen daraus, vorbehalten. Kein Teil des Werkes darf ohne schriftliche Genehmigung des Verlages in irgendeiner Form reproduziert oder unter Verwendung elektronischer Systeme verarbeitet, vervielfältigt oder verbreitet werden. Die Wiedergabe von Gebrauchsnamen, Warenbezeichnungen, usw. in diesem Werk berechtigt auch ohne besondere Kennzeichnung nicht zu der Annahme, dass solche Namen im Sinne der Warenzeichen- und Markenschutz-Gesetzgebung als frei zu betrachten wären und daher von jedermann benutzt werden dürfen. This document is protected by copyright law. No part of this document may be reproduced in any form by any means without prior written authorization of the publisher. Alle Rechte vorbehalten | all rights reserved © Mensch und Buch Verlag 2014 Choriner Str. 85 - 10119 Berlin [email protected] – www.menschundbuch.de To my mother Gisela, my father Martin, my brother Nikolas and my aunt Barbara for their endless love and unmeasurable support Der Tod ist kein Abschnitt des Daseins, sondern nur ein Zwischenereignis, ein Übergang aus einer Form des endlichen Wesens in eine andere. (Wilhelm von Humboldt, Briefe an eine Freundin) Contents I. Introduction ......................................................................................................................... 9 1. Embryonic resorption in mammals ................................................................................... 9 1.1. Historical overview ..................................................................................................... 9 1.2. Definition .................................................................................................................. 11 1.3. Embryonic resorption in polytocous species ............................................................ 11 1.3.1. Macroscopic description ................................................................................ 11 1.3.2. Incidence ........................................................................................................ 12 1.3.2.1. Direct counts of embryonic resorptions ....................................... 12 1.3.2.2. The reliability of corpora lutea counts ......................................... 13 1.3.2.3. The reliability of placental scar counts ........................................ 13 1.4. Embryonic degeneration in monotocous species..................................................... 14 1.4.1. Incidence ........................................................................................................ 14 1.4.1. Multiple pregnancies ...................................................................................... 15 2. The European brown hare .............................................................................................. 15 2.1. Population decline .................................................................................................... 16 2.2. Factors associated with polpulation declines in Europe ........................................... 16 2.3. Reproductive research ............................................................................................. 17 2.4. Incidence of embryonic resorption in the European brown hare .............................. 18 3. Biopsy procedures and Amniocentesis ........................................................................... 18 3.1. Amniocentesis .......................................................................................................... 18 3.2. Chorionic villus sampling ......................................................................................... 19 3.3. Ultrasound guided assistance .................................................................................. 19 4. Objectives ....................................................................................................................... 20 II. Articles .............................................................................................................................. 21 Article I. Embryonic resorption in context to intragestational corpus luteum regression: a longitudinal ultrasonographic study in the Eurpean brown hare (Lepus europaeus PALLAS 1778). ..... 21 Article II. In vivo tissue sampling of embryonic resorption sites using ultrasound guided biopsy.. .... 37 Article III. Free blastocyst and implantation stages in the European brown hare: correlation between ultrasound and histological data.. ....................................................................................... 52 III. General discussion ......................................................................................................... 76 IV. Conclusion: Embryonic resorption as a physiological phenomenon in mammals.. 79 V. Summary/Zusammenfassung ......................................................................................... 84 VI. References ....................................................................................................................... 86 VII. Appendix ....................................................................................................................... 116 5 This thesis is based on the following manuscripts: 1. Schroeder, K., Drews, B., Roellig, K., Goeritz, F. and Hildebrandt, T.B. (2013) ‘Embryonic resorption in context to intragestational corpus luteum regression: a longitudinal ultrasonographic study in the European brown hare (Lepus europaeus PALLAS 1778)’, Theriogenology, vol. 80, pp. 778-784. - own contributions: ultrasonographic evaluation from day 5 to day 42 of pregnancy of 1) embryonic development, 2) embryonic resorptions and 3) CL-monitoring, 4) analysis and statistical calculations, 5) writing of the manuscript 2. Schroeder, K., Drews, B., Roellig, K., Menzies, B.R., Goeritz, F. and Hildebrandt, T.B. (2011) ‘In vivo tissue sampling of embryonic resorption sites using ultrasound guided biopsy’, Theriogenology, vol. 76, pp. 778-784. - own contributions: 1) implementation of the two biopsies techniques and in vivo extraction of resorption fluid and placental tissue, 2) ultrasonographic monitoring of embryonic resorptions and viable conceptus from day 5 to day 42, 3) histological analysis of resorption smears, 4) analysis and statistical calculations, 5) writing of the manuscript 3. Drews, B., Ringleb, J., Waurich, R., Hildebrandt, T.B., Schröder, K. and Roellig, K. (2013) ‘Free blastocyst and implantation stages in the European brown hare: correlation between ultrasound and histological data’, Reproduction, Fertility and Development, vol. 25, pp. 866-878. – own contributions: 1) ultrasonographic monitoring and 2) tissue collection 6 List of abbreviations: AES agri-environment schemes AI artificial insemination BVDV bovine viral diarrhea virus CL corpus luteum, corpora lutea CS caesarean section CT computed tomography CVS chorionic villus sampling d day DNA desoxyribonucleic acid Ebh European brown hare EBHS European Brown Hare Syndrome eCG equine chorionic gonadotropin EED early embryonic death eFh Europäischer Feldhase em embryo EPL early pregnancy loss ET embryo transfer GK Gelbkörper h hour kg kilogram LAP laparatomy LH luteinizing hormone mg milligram Mhz Megahertz min minute ml millilitre MRI magnetic resonance imaging ng nanogram OCT optical coherence tomography OHE ovariohysterectomy pl placental PLR partial litter resorption p.m. post mortem RES resorption s.d. standard deviation SRY sex determining region of Y 7 TLR total litter resorption UBM ultrasound biomicroscope US ultrasound µg microgram µl microlitre µm micrometre µM micromole 8 I. Introduction 1. Embryonic resorption in mammals 1.1. Historical overview The phenomenon of embryonic resorption
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