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Common Principles of Olfactory Coding Across Olfactory Receptor Families and Species

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Common Principles of Olfactory Coding Across Olfactory Receptor Families and Species Common principles of olfactory coding across olfactory receptor families and species Inauguraldissertation zur Erlangung des Doktorgrades der Mathematisch-Naturwissenschaftlichen Fakultät der Universität zu Köln vorgelegt von Daniel Kowatschew aus Hoyerswerda (Copy Star,Köln) 2021 Berichterstatter/in: Prof. Dr. Sigrun Korsching und Prof. Dr. Kay Hofmann Tag der letzten mündlichen Prüfung: 19.02.2021 I Abstract Beside vision and hearing, the chemoreception is one of the important senses for organ- isms in an aquatic environment. To detect food, predators or sexual partners a broad vari- ation of different receptors in the nose of an animal is needed. Therefore, coding strategy of the olfactory receptors is the formation of spatial distribution patterns in order to maintain their continued function in the event of damage to the epithelium. The investiga- tion of individual receptor groups of the olfactory system in fish is now mostly done through phylogenetic studies, as more and more genomes are being deciphered enabled by rapid technical progress. This work focused on the visualization of individual receptors and re- ceptor families of different aquatic animals in the olfactory epithelium and for this, among other things, expression and spatial distribution were investigated. Here it could be shown, that the recently discovered adenosine receptor A2c is expressed in the eel, carp and claw frog in a sparse and distributed pattern within their olfactory epithelium similar to the pat- tern observed for zebrafish. A sharp contrast is formed by the expression of A2c in lamprey, because of A2c-expressing cells forming a contiguous domain directly adjacent to the sensory region. In addition it was shown, that this expression is consistent, with the expression in neuronal progenitor cells. It could be that A2c switched the function during evolution in the lineage of vertebrates. Also for the first time, the expression pattern of three of six possible V1R receptors, and the absence of two possible V1R and two V2R re- ceptors in adult Lampetra fluviatilis was demonstrated. It was shown that the three V1Rs form different expression zones, when looking at the height in the lamella. Here we also completed a project to investigate the spatial distribution of the complete V1R-related Oras receptor family in zebrafish. It was shown, that each of the ora genes had its own expres- sion zone. Beside this, two different in situ staining methods were used and compared with each other, chromogenic NBT/BCIP staining and fluorescent TSA staining. It turned out that the TSA method is the better choice, as it is much more sensitive, and only through this was it possible to examine the expression pattern of two of the six V1RS, Ora5 and Ora6. II Zusammenfassung Neben dem Sehen und Hören ist die Chemorezeption einer der wichtigsten Sinne für Or- ganismen in Gewässern. Um Nahrung, Raubtiere oder Sexualpartner zu erkennen, ist eine breite Variation verschiedener Rezeptoren in der Nase des Tieres erforderlich. Die Kodier- ungsstrategie der Geruchsrezeptoren ist die Bildung räumlicher Verteilungsmuster, um ihre Funktion im Falle einer Schädigung des Epithels aufrechtzuerhalten. Die Unter- suchung einzelner Rezeptorgruppen des olfaktorischen Systems in Fischen erfolgt heute hauptsächlich durch phylogenetische Studien, da aufgrund des raschen technischen Fortschritts immer mehr Genome entschlüsselt werden. Diese Arbeit konzentrierte sich auf die Visualisierung einzelner Rezeptoren und Rezeptorfamilien verschiedener Wassertiere im Riechepithel und untersuchte unter anderem deren Expression und räumliche Ver- teilung. Hier konnte gezeigt werden, dass der kürzlich entdeckte Adenosinrezeptor A2c in Aal, Karpfen und Klauenfrosch in einem spärlich verteilten Muster innerhalb ihres Riechepithels exprimiert wird, welches dem für Zebrafische beobachteten Muster sehr ähnlich ist. Ein scharfer Kontrast dazu, ist das Expressionsmuster von A2c im Neunauge, da die A2c-positiven Zellen dort eine zusammenhängende Domäne direkt neben der sen- sorischen Region bilden. Zusätzlich aufgezeigt wurde, dass die Lokalisation der A2c Rezeptoren mit der Expressionsregion von neuronalen Vorläuferzellen übereinstimmt. De- shalb könnte es sein, dass A2c die Funktion während der Evolution in der Linie der Wirbel- tiere wechselte. Des Weiteren konnte hier zum ersten Mal das Expressionsmuster von drei möglichen V1R-Rezeptoren und das Fehlen dieses bei zwei möglichen V1R- und zwei V2R-Rezeptoren bei adulten Neunaugen (Lampetra fluviatilis) bestätigt werden. Bei der Untersuchung der Höhe in der Lamelle, wurde beobachtet, dass die drei V1Rs ver- schiedene Expressionszonen bilden. Zusätzlich konnte ein Projekt welches die Unter- suchung der räumlichen Verteilung der gesamten V1R-verbundenen Ora Familie im Zebrafisch beinhaltete abgeschlossen werden. Es konnte gezeigt werden, dass jedes der ora-Gene eine eigene Expressionszone besitzt. Dafür wurden zwei verschiedene in situ -Färbemethoden verwendet und miteinander verglichen, die chromogene NBT/BCIP-Fär- bung und die fluoreszierende TSA-Färbung. Es stellte sich dabei die TSA-Methode als die bessere Wahl heraus, da diese viel sensitiver ist, und nur dadurch das Expressionsmuster von zwei der sechs V1RS, Ora5 und Ora6, untersucht werden konnte. III Table of content 1.INTRODUCTION...............................................................................................................1 1.1 Purines and Purinergic receptors...............................................................................1 1.2 G protein-coupled receptor.........................................................................................5 1.3 Olfactory receptors.....................................................................................................8 1.3.1 ORs.....................................................................................................................9 1.3.2 V1Rs.................................................................................................................10 1.3.3 V2Rs.................................................................................................................11 1.3.4 TAARs...............................................................................................................11 1.3.5 FPRs.................................................................................................................12 1.3.6 GUCYs..............................................................................................................12 1.3.7 MS4As..............................................................................................................13 1.3.8 A2C...................................................................................................................14 1.4 Olfactory systems.....................................................................................................14 1.4.1 The olfactory system of zebrafish (Danio).........................................................14 1.4.2 The olfactory system of the eel (Anguilla).........................................................16 1.4.3 The olfactory system of carp (Cyprinus) ..........................................................17 1.4.4 The olfactory system of the claw frog (Xenopus)..............................................18 1.4.5 The olfactory system of lamprey.......................................................................19 2. Material and Methods ....................................................................................................23 2.1 Technical equipment................................................................................................23 2.2 Chemicals................................................................................................................24 2.2.1 Solutions and buffers for long term storage ….................................................. 24 2.2.2 Short term storage solutions and buffers..........................................................25 2.3 Antibody...................................................................................................................27 2.4 Primer.......................................................................................................................28 2.5 Molecular biology techniques...................................................................................32 2.5.1 Synthesis of Digoxigenin and Biotin labelled Thyramides.................................32 2.5.2 Production of cDNA ..........................................................................................32 2.5.3 DNA extraction from tissue ..............................................................................33 2.5.4 Fin clipping and fast DNA extraction for zebrafish............................................33 IV 2.5.5 Production of molecular probes .......................................................................34 2.5.5.1 PCR...........................................................................................................34 2.5.5.2 Ligation of the PCR products into pGEM-T vectors...................................35 2.5.5.3 The making of the agar plates...................................................................35 2.5.5.4 Transforming the plasmid DNA into electrocompetent cells......................35 2.5.5.5 Picking the colonies...................................................................................36 2.5.5.6 Plasmid purification....................................................................................36
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