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Diplozoidae, Monogenea) – an Analysis of Selected Organ Systems

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Diplozoidae, Monogenea) – an Analysis of Selected Organ Systems MASARYK UNIVERSITY FACULTY OF SCIENCE DEPARTMENT OF BOTANY AND ZOOLOGY Ultrastructural studies on diplozoid species (Diplozoidae, Monogenea) – an analysis of selected organ systems Ph.D. Dissertation Veronika Konstanzová Supervisor: Assoc. Prof. RNDr. Milan Gelnar, CSc. BRNO 2017 Bibliographic Entry Author: Mgr. Veronika Konstanzová Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Ultrastructural studies on diplozoid species (Diplozoidae, Monogenea) - an analysis of selected organ systems Degree programme: Biology Field of Study: Parasitology Supervisor: Assoc. Prof. RNDr. Milan Gelnar, CSc. Faculty of Science, Masaryk University Department of Botany and Zoology Academic Year: 2016/2017 Number of Pages: 135 Keywords: Monogenea; Diplozoid species; Morphology; Ultrastructure; Gastro-intestinal tract; Excretory system; Neodermis; Clamps Bibliografický záznam Autor: Mgr. Veronika Konstanzová Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Ultrastrukturní studie zástupců čeledi Diplozoidae (Monogenea) – analýza vybraných orgánových soustav Studijní program: Biologie Studijní obor: Parazitologie Školitel: Doc. RNDr. Milan Gelnar, CSc. Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Akademický rok: 2016/2017 Počet stran: 135 Klíčová slova: Monogenea; Diplozoidae; Morfologie; Ultrastruktura; Trávicí soustava; Exkreční soustava; Neodermis, Svorky ABSTRACT Diplozoids are representatives of blood-feeding ectoparasites from the family Diplozoidae (Polyopisthocotylea, Monogenea). Although these helminths have been the subject of numerous taxonomical, phylogenetic and ecological studies, information about their organ morphology and ultrastructure remain mostly unknown. The Ph.D. dissertation is composed of three studies, each dealing with a specific topic: A) An ultrastructural analysis of gastro-intestinal tract of Paradiplozoon homoion. B) An ultrastructural analysis of excretory system of four diplozoid species: Paradiplozoon bliccae, Paradiplozoon homoion, Diplozoon paradoxum and Eudiplozoon nipponicum. C) An ultrastructural analysis of neodermal surface and haptoral attachment clamps of Paradiplozoon homoion. Diplozoids were obtained from the gills of several species of cyprinid fish. The worms used for ultrastructural studies were fixed by osmium tetroxide, dehydrated through ascending acetone series and embedded in a Spurr resin. Ultrathin sections were contrasted and examined by using a transmission electron microscopy (TEM). Paper A - the observation on gastrointestinal tract of P. homoion revealed two main types of digestive cells — U-shaped digestive cells and connecting syncytium. Particular structures such as mouth cavity, two oval-shaped muscular buccal suckers, pharynx surrounded with the glandular cells, oesophagus, the intestinal caeca with intact erythrocytes of the host fish in the lumen, the apical pinocytotic fibrous surface complex and haematin vesicles of U-shaped cells have been shown in detail. Paper B – ultrastructure of two basic compartments which forms the excretory apparatus, flame cells with filtration apparatus, and canal cells forming the protonephridial ducts were revealed. A unique sequence of semi-thin sections of the excretory pore of E. nipponicum lined by tegument invaginated from the neodermal surface layer was documented there for the first time. Paper C - provides detailed ultrastructural data on the neodermal external layer and compartments of haptoral attachment clamps of P. homoion. Glandular cells at the base of the attachment clamps and a presence of electron-dense material responsible for sclerite development in diplozoid species were observed for the first time in monogeneans. ABSTRAKT Diplozooni jsou zástupci krevsajících ektoparazitů z čeledi Diplozoidae (Polyopisthocotylea, Monogenea). Přestože byli tito helminti předmětem četných výzkumů zaměřených na studium taxonomie, fylogenetiky a ekologických studií, informace o vnitřní struktuře jejich těla a ultrastruktuře orgánů chybí. Předložená dizertační práce se skládá ze tří studií zaměřených na následující témata: A) Analýza ultrastruktury trávicího traktu druhu Paradiplozoon homoion. B) Analýza ultrastruktury vylučovacího systému u čtyř různých druhů diplozoonů: Paradiplozoon bliccae, Paradiplozoon homoion, Diplozoon paradoxum a Eudiplozoon nipponicum. C) Analýza ultrastruktury povrchu těla (neodermis) a svorek příchytného aparátu druhu Paradiplozoon homoion. Zástupci čeledi Diplozoidae byli získáni ze žáber několika druhů kaprovitých ryb. Jedinci vybraní pro ultrastrukturální analýzy byli fixováni oxidem osmičelým, odvodněni pomocí acetonové vzestupné řady a zaliti do pryskyřice Spurr. Ultratenké řezy byly kontrastovány a prohlíženy v transmisním elektronovém mikroskopu (TEM). Práce A - studium trávicího traktu druhu P. homoion prokázalo dva hlavní typy buněk, z nichž se skládá střevo diplozoona – trávicí buňky tvaru U a podpůrné spojovací soubuní. Dále byly detailně studovány další části trávicího traktu, jako je ústní dutina, dvě oválné svalnaté bukální přísavky, hltan obklopený žláznatými buňkami, jícen, střevo s nepoškozenými červenými krvinkami hostitelské ryby, povrch trávicích buněk s vláknitými strukturami. Práce B analyzuje ultrastrukturu vylučovací soustavy, zejména jejích dvou základních stavebních jednotek – plaménkových buněk a kanálkových buněk, které jsou stavební jednotkou vylučovacích kanálků. V této práci je poprvé u monogeneí publikována série mikrofotografií řezů exkrečním pórem druhu E. nipponicum. Exkreční pór je vystlán povrchovou vrstvou neodermis. Práce C je zaměřena na detailní ultrastrukturální analýzu neodermálního povrchu těla diplozoonů a jednotlivých komponent příchytných svorek haptoru u druhu P. homoion. U zástupců čeledi Diplozoidae byl v této práci publikován výskyt žláznatých buněk u báze příchytných svorek haptoru a také přítomnost elektron-denzního materiálu, který je pravděpodobně zodpovědný za růst a rozvoj skleritů tvořících kostru svorky. © Veronika Konstanzová, Masaryk University, 2017 “A picture speaks a thousand words…” Javier del Campo et al. (2016) ACKNOWLEDGEMENTS I would like to express my sincere thanks to my supervisor Associate Professor Milan Gelnar for his guidance, comprehensive help, the opportunity to work in his laboratory and for financial support. Great thanks go to Dr. Božena Koubková, who genuinely helped me during my whole studies with all the work that finally brought this thesis into being. I would also like to thank Professor Iva Dyková for support with TEM observations, helpful comments and kind approach during my Ph.D. study. My sincere thanks belong to all the colleagues that helped me in the laboratory or in any other way and to Kevin Roche for English correction of the dissertation. Many thanks also go to my husband Roman and my family for supporting me in all possible ways. TABLE OF CONTENTS 1. INTRODUCTION…………………………………...……………………... 14 2. AIMS OF THE STUDY……………………………..................................... 16 3. LITERATURE OVERVIEW……………………………………………… 17 3.1 Family Diplozoidae…………………………………..................................... 17 3.1.1 Taxonomic history of Diplozoidae, Palombi, 1949………………………... 17 3.1.2 Life cycle of Diplozoinae, Palombi, 1949…..……………………………... 19 3.1.3 Localization of diplozoids on gills of fish host……..................................... 20 3.1.4 Opisthaptor attachment apparatus……………............................................. 21 3.2 Morphology and ultrastructure of selected blood-feeding polyopisthocotylean organ systems ………………………………………. 23 3.2.1 Gastro-intestinal system...…………………………………………………. 24 3.2.2 Excretory system…………………………………………………………... 25 3.2.3 Surface (neodermis)………………………………………………………... 27 3.2.4 Attachment apparatus……………………………………………………… 28 4. MATERIAL AND METHODS…………………………………………… 30 4.1 Species under study, hosts and locality…………........................................ 30 4.2 Transmission electron microscopy (TEM)……........................................... 31 5. SUMMARY OF RESULTS…………………………………………………. 33 5.1 Paper A……………………………………………………………………… 33 5.2 Paper B……………………………………………………………………… 35 5.3 Paper C……………………………………………………………………… 36 6. DISCUSSION……………………………………............................................ 39 7. CONCLUSIONS AND PERSPECTIVE…………….…………...………… 46 8. REFERENCES……………………………………………............................. 48 9. PUBLICATIONS FORMING THE DISSERTATION…………………… 67 10. SUPPLEMENTARY MATERIAL………………………………………... 99 10.1 Gastro-intestinal system…………………………………………………... 99 10.2 Excretory system………………………………………………………… 104 10.3 Surface (neodermis)……………………………………………………... 110 10.4 Attachment apparatus…………………………………………………... 112 10.5 Reproductive system…………………………………………………….. 113 11. APPENDIX………………………………………………………………... 119 11.1 Author contributions to the papers presented in the thesis…………... 119 11.2 Other publication unrelated to the dissertation……………………….. 121 1 INTRODUCTION Diplozoids (Polyopisthocotylea, Heteronchoinea) are blood-feeding monogeneans, mainly parasitising the gills of cyprinid fish, that are characterised by a unique X-shaped body arrangement, the result of two adult individuals permanently fusing together (Fig. 1). The adult worms represent a health risk in fish aquaculture, where they cause diseases problems such as anaemia (Kawatsu 1978) and weight loss (Anderson 1974). Impacts on fish fitness often result in a decrease in natural body resistance, with consequent development of secondary microbial infections (Buchman and Bresciani
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