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Ontogenetska, Histološka, Semiohemijska I Etološka Istraživanja Vrste Pachyiulus Hungaricus (Karsch, 1881) (Diplopoda, Julida, Julidae)

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Ontogenetska, Histološka, Semiohemijska I Etološka Istraživanja Vrste Pachyiulus Hungaricus (Karsch, 1881) (Diplopoda, Julida, Julidae) UNIVERZITET U BEOGRADU BIOLOŠKI FAKULTET Zvezdana S. Jovanović ONTOGENETSKA, HISTOLOŠKA, SEMIOHEMIJSKA I ETOLOŠKA ISTRAŽIVANJA VRSTE PACHYIULUS HUNGARICUS (KARSCH, 1881) (DIPLOPODA, JULIDA, JULIDAE) doktorska disertacija Beograd, 2019 UNIVERSITY OF BELGRADE FACULTY OF BIOLOGY Zvezdana S. Jovanović ONTOGENETIC, HISTOLOGICAL, SEMIOCHEMICAL AND ETHOLOGICAL RESEARCH OF PACHYIULUS HUNGARICUS (KARSCH, 1881) (DIPLOPODA, JULIDA, JULIDAE) Doctoral Dissertation Belgrade, 2019 Mentori: ____________________________________ Prof. dr Slobodan Makarov, redovni profesor, Univerzitet u Beogradu - Biološki fakultet ____________________________________ Prof. dr Luka Lučić, redovni profesor, Univerzitet u Beogradu - Biološki fakultet Članovi komisije: ____________________________________ Prof. dr Aleksandra Korać, redovni profesor Univerzitet u Beogradu - Biološki fakultet ____________________________________ Prof. dr Sofija Pavković-Lučić, redovni profesor Univerzitet u Beogradu - Biološki fakultet ____________________________________ Dr Ljubodrag Vujisić, docent Univerzitet u Beogradu - Hemijski fakultet Datum odbrane: ______________ ZAHVALNICA Ova doktorska disertacija je urađena na Katedri za dinamiku razvića životinja Univerziteta u Beogradu - Biološkog fakulteta, i predstavlja rezultat saradnje sa Katedrom za biologiju ćelija i tkiva i Centrom za elektronsku mikroskopiju, Katedrom za mikrobiologiju Univerziteta u Beogradu - Biološkog fakulteta i Katedrom za organsku hemiju Univerziteta u Beogradu - Hemijskog fakulteta. Finansijska podrška obezbeđena je projektom Ministarstva prosvete, nauke i tehnološkog razvoja pod nazivom “Ontogenetska karakterizacija filogenije bioraznovrsnosti” (OI 173038). S obzirom da su u ovoj tezi prikazani rezultati istraživanja različitih aspekata biologije određenog model organizma, zahvalnost dugujem mnogobrojnim profesorima, saradnicima i istraživačima, a pre svega svojim mentorima, redovnim profesorima dr Slobodanu Makarovu i dr Luki Lučiću sa Katedre za dinamiku razvića životinja, kao i članovima komisije, prof. dr Aleksandri Korać sa Katedre za biologiju ćelija i tkiva, prof. dr Sofiji Pavković-Lučić sa Katedre za genetiku i evoluciju i doc. dr Ljubodragu Vujisiću sa Katedre za organsku hemiju Univerziteta u Beogradu - Hemijskog fakulteta. Zahvaljujem se naučnom saradniku dr Bojanu Iliću i ostalim članovima Katedre za dinamiku razvića životinja, naučnom saradniku dr Vukici Vujić sa Katedre za genetiku i evoluciju, zatim članovima Katedre za biologiju ćelija i tkiva i Centra za elektronsku mikroskopiju, Katedre za mikrobiologiju Univerziteta u Beogradu - Biološkog fakulteta, Katedre za organsku hemiju Univerziteta u Beogradu - Hemijskog fakulteta, i prof. dr Draganu Nikoliću pri Laboratoriji za elektronsku mikroskopiju Univerziteta u Beogradu - Poljoprivrednog fakulteta. Takođe, hvala mojim koleginicama sa doktorskih studija Mariji Aleksić i Ivani Šaganović. Hvala mojoj mami, ostalim članovima porodice i Nemanji Jovanoviću. Fibonačijev niz: Pachyiulus hungaricus (Karsch, 1881) u odbrambenom položaju Ontogenetska, histološka, semiohemijska i etološka istraživanja vrste Pachyiulus hungaricus (Karsch, 1881) (Diplopoda, Julida, Julidae) Sažetak U ovoj tezi predstavljeni su rezultati istraživanja različitih aspekata biologije diplopode Pachyiulus hungaricus (Karsch, 1881): kasnog perioda postembrionalnog razvića, histološke građe odbrambenih žlezda, sastava i antimikrobnog dejstva odbrambenog sekreta i reproduktivnog ponašanja. Euanamorfno razviće je proučavano kroz stadijume VII-XII, odnosno, kroz period kasne postembriogeneze. Definisanje stadijuma izvedeno je na osnovu broja nizova ocela, dok vrednosti analiziranih morfoloških osobina imaju dopunski informacioni značaj. Adultne jedinke sa funkcionalnim gonopodama i funkcionalnim vulvama uočene su na stadijumu XI. Histološka građa odbrambenih žlezda je analizirana kod adultnih jedinki. Komponente odbrambenog sekreta su lokalizovane u sekretornim granulama, koje se, na osnovu određenih morfoloških osobina mogu svrstati u nekoliko tipova. Rezultati ultrastrukturne analize ukazuju da se sekret kanalima doprema u lumen rezervoara, što odgovara merokrinom tipu sekrecije. Dominantna jedinjenja u sekretu P. hungaricus su hinoni, dok su nehinonska jedinjenja u vidu pentil i heksil estara zasićenih i nezasićenih masnih kiselina dugog niza. Sekret je ispoljio inhibitorno dejstvo na rast gljiva i bakterija, kao i baktericidno i fungicidno dejstvo. S obzirom da u privlačenju partnera mogu da deluju i hemijski stimulusi, proučavan je aspekat reproduktivnog ponašanja, prekopulatorno ponašanje. Opisano je nekoliko sekvenci ponašanja u testu nazvanom „arena za parenje“, a zatim u testovima izbora mužjaka i izbora ženki. Nekoliko morfoloških osobina jedinki koje su se parile i onih koje nisu imale uspeha u parenju, analizirano je metodama tradicionalne i geometrijske morfometrije. Rezultati su pokazali da je osobina oblika nogu podložna prekopulatornoj seksualnoj selekciji. Ključne reči: Pachyiulus hungaricus, Diplopoda, euanamorfoza, morfološka varijabilnost, odbrambene žlezde, sekretorne ćelije, odbrambeni sekret, antimikrobno dejstvo, uspeh u parenju. Naučna oblast: Biologija Uža naučna oblast: Biologija razvića životinja UDK broj: [591.8+591.572.5+591.351]: 595.61(043.3) Ontogenetic, histological, semiochemical and ethological researchof species Pachyiulus hungaricus (Karsch, 1881) (Diplopoda, Julida, Julidae) Abstract In this thesis, the results of research on various aspects of biology of diplopod Pachyiulus hungaricus (Karsch, 1881) are presented: late period of postembryonic development, histological structure of defensive glands, composition and antimicrobial effect of defensive secretion and reproductive behaviour. Stadia VII-XII, which belong to the late period of euanamorphic development, were included in the analysis. Counting rows of ocelli was the primary method for differentiating between stadia, while the other analysed morphological traits are of supplementary informational significance. Adults possess functional gonopods and functional vulvae at the stadium XI. Histological structure of defensive glands was investigated in adults. Components of defensive secretion are localized in secretory granules, which could be classified into several types, based on their morphology. Results of ultrastructural analysis indicate that ducts transport the secretion into reservoir lumen, which corresponds to the merocrine secretion mode. Dominant compounds in the secretion of P. hungaricus are quinones, and non- quinonic compounds are pentyl and hexyl esters of long-chain fatty acids. The secretion had inhibitory effect on the growth of tested bacteria and fungi, as well as bactericidal and fungicidal effect. Considering that chemical stimuli could be important in the mate attraction, we studied the aspect of reproductive behaviour, precopulatory behaviour. Several sequences of the behaviour were described during mating arena test and then in male and female choice tests. Also, several morphological traits of mated and nonmated individuals were analysed using traditional and geometric morphometrics. The results indicate that the leg shape is subjected to precopulatory sexual selection. Key words: Pachyiulus hungaricus, Diplopoda, euanamorphosis, morphological variability defensive glands, secretory cells, defensive secretion, antimicrobial effect, mating success. Scientific field: Biology Scientific subfield: Biology of Animal Development UDC number: [591.8+591.572.5+591.351]: 595.61(043.3) Sadržaj 1. UVOD 1 1.1. Opšte odlike klase Diplopoda 1 1.2. Postembriogeneza Diplopoda 6 1.2.1 Anamorfno razviće Diplopoda 6 1.2.2 Definisanje postembrionalnih stadijuma Diplopoda 12 1.3. Anatomske i fiziološke odlike odbrambenih žlezda Diplopoda 13 1.4. Odbrambeni mehanizmi klase Diplopoda 16 1.4.1. Hemijska odbrana 20 1.4.1.1. Glomeridni tip žlezda 21 1.4.1.2. Polidezmidni tip žlezda 22 1.4.1.3. Julidni tip žlezda 25 1.5. Poliginandrija, seksualna selekcija i 27 reproduktivno ponašanje Diplopoda 2. CILJEVI ISTRAŽIVANJA 31 3. MATERIJAL I METODE 32 3.1. Pachyiulus hungaricus – rasprostranjenje i osnovne morfološke odlike 32 3.2. Prikupljanje i čuvanje jedinki u laboratorijskim uslovima 33 3.3. Metode istraživanja postembrionalnog razvića 34 3.3.1. Prikupljanje i merenje jedinki 34 3.3.2. Analizirane morfološke osobine 34 3.3.3. Metode određivanja postembrionalnih stadijuma P. hungaricus 35 3.4. Metode histoloških istraživanja 36 3.4.1. Priprema odbrambenih žlezda za elektronsku mikroskopiju 36 3.4.2. Bojenje toluidinom 37 3.4.3. Imerzija ozadena u mlečnu kiselinu 37 3.4.4. Merenje morfoloških celina, histoloških i 38 citoloških komponenti 3.5. Metode semiohemijskih istraživanja 38 3.5.1. Prikupljanje i priprema odbrambenog sekreta 38 3.5.2. Hemijske analize 38 3.5.3. Patogeni sojevi i uslovi gajenja kultura 39 3.5.4. Antibakterijska aktivnost odbrambenog sekreta P. hungaricus 40 3.5.5. Antifungalna aktivnost odbrambenog sekreta P. hungaricus 42 3.6. Metode etoloških istraživanja 42 3.6.1. Uzorkovanje i pripremanje jedinki za testove parenja 42 3.6.2. Testovi parenja 43 3.6.3. Analiza odabranih morfoloških osobina kod P. hungaricus 44 4. REZULTATI 47 4.1. Rezultati istraživanja postembrionalnog razvića 47 4.1.2. Postembrionalni stadijumi P. hungaricus 47 4.2. Rezultati histoloških istraživanja 53 4.2.1. Morfološke i histološke odlike odbrambenih žlezda P. hungaricus 53 4.2.2. Ultrastrukturna analiza odbrambenih žlezda P. hungaricus 58 4.3. Rezultati semiohemijskih istraživanja 65 4.3.1.
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