Uporedni pregled morfo-anatomskih karakteristika biljnih organa i analiza etarskih ulja sa njihovom primenom u taksonomiji odabranih rodova tribusa Inuleae Cass. (Compositae) / Comparative review of morpho-anatomical characteristics of plant organs and analyses of essential oils with their application in taxonomy of selected genera of the tribe Inuleae Cass. (Compositae)

Description

<p>Komparativnom&nbsp; analizom&nbsp; morfo-anatomskih&nbsp; i mikromorfolo&scaron;kih&nbsp; karakteristika&nbsp; lista,&nbsp; stabla,&nbsp; rizoma,&nbsp; korena, ploda&nbsp; i&nbsp; receptakuluma,&nbsp; kao&nbsp; i&nbsp; biohemijskom&nbsp; analizom&nbsp; etarskih ulja&nbsp; podzemnih&nbsp; organa,&nbsp; obuhvaćeno&nbsp; je&nbsp; ukupno&nbsp; 16&nbsp; vrsta&nbsp; iz&nbsp; 4<br />roda,&nbsp; Inula&nbsp; (I. aschersoniana,&nbsp; I. bifrons,&nbsp; I. britannica,&nbsp;<em> I. conyza, I.&nbsp; ensifolia,&nbsp; I.&nbsp; germanica,&nbsp; I.&nbsp; helenium,&nbsp; I.&nbsp; oculus-christi,&nbsp; I. saslicina&nbsp; ssp.&nbsp; salicina,&nbsp; I.&nbsp; salicina&nbsp; ssp.&nbsp; aspera,&nbsp; I.&nbsp; spiraeifolia), Dittrichia&nbsp; (D.&nbsp; graveolens,&nbsp; D.&nbsp; viscosa),&nbsp; Limbarda&nbsp; (L.<br />crithmoides)&nbsp; i&nbsp; Pulicaria&nbsp; (P. dysenterica,&nbsp; P. vulgaris) iz</em> tribusa Inuleae,&nbsp; familije&nbsp; Compositae.&nbsp; Vrste&nbsp; iz&nbsp; različitih&nbsp; rodova analizirane&nbsp; u&nbsp; ovoj&nbsp; disertaciji&nbsp; ranijom&nbsp; klasifikacijom&nbsp; svrstavane su u okviru različitih sekcija roda&nbsp; Inula, a takođe i tipska vrsta ovog roda,<em> I. helenium L</em>., svrstavana je u okviru zasebne sekcije<em> Corvisartia.</em>&nbsp; Na&nbsp; osnovu&nbsp; najnovijih&nbsp; molekularnih&nbsp; filogenetskih analiza, baziranih na sekvencama plastidne i jedarne DNK, kao i na osnovu morfolo&scaron;kih i kariolo&scaron;kih podataka, predloženo je da sekcija&nbsp; <em>Corvisartia&nbsp;</em> bude&nbsp; izdvojena&nbsp; u&nbsp; zaseban&nbsp; rod.&nbsp; Ciljevi&nbsp; ove<br />disertacije definisani su u skladu sa činjenicom da su podaci o morfo-anatomskoj,&nbsp; mikromorfolo&scaron;koj&nbsp; i&nbsp; biohemijskoj&nbsp; građi vegetativnih&nbsp; i&nbsp; reproduktivnih&nbsp; organa&nbsp; vrsta&nbsp; ovog&nbsp; tribusa malobrojni&nbsp; i&nbsp; nepotpuni,&nbsp; dok&nbsp; su&nbsp; analize&nbsp; vr&scaron;ene&nbsp; parcijalno&nbsp; i<br />nezavisno&nbsp; kod&nbsp; pojedinih&nbsp; predstavnika,&nbsp; sa&nbsp; slabim&nbsp; akcentom&nbsp; na njihovu&nbsp; komparativnu&nbsp; analizu.&nbsp; Istraživanje&nbsp; je&nbsp; obuhvatilo&nbsp; veliki broj kvalitativnih i kvantitativnih karaktera koji su analizirani i mereni&nbsp; pomoću&nbsp; stereomikroskopa,&nbsp; svetlosnog,&nbsp; fluorescentnog&nbsp; i skening elektronskog mikroskopa. Dobijeni podaci statistički su obrađeni&nbsp; različitim&nbsp; metodama&nbsp; numeričke&nbsp; analize&nbsp; koje&nbsp; se&nbsp; inače<br />koriste pri taksonomskim istraživanjima.&nbsp; Ovakav tip analize za vrste&nbsp; rodova&nbsp;<em> Inula,</em>&nbsp; <em>Dittrichia,&nbsp; Limbarda&nbsp; i&nbsp; Pulicaria</em>&nbsp; nije&nbsp; zabeležen u dosada&scaron;njoj literaturi.<br />Sprovedena&nbsp; analiza&nbsp; pokazala&nbsp; je&nbsp; da&nbsp; ni&nbsp; jedan&nbsp; vegetativni&nbsp; biljni organ&nbsp; pojedinačno&nbsp; ne&nbsp; pruža&nbsp; dovoljno&nbsp; informacija&nbsp; na&nbsp; osnovukojih bi se sa statističkom sigurno&scaron;ću mogle izdvojiti određene grupe&nbsp; vrsta,&nbsp; niti&nbsp; rodovi&nbsp; prema&nbsp; postojećoj&nbsp; podeli.&nbsp; U&nbsp; okviru analiziranih anatomskih i mikromorfolo&scaron;kih karakteristika ploda i&nbsp; receptakuluma,&nbsp; organizacija&nbsp; sklerenhimskog&nbsp; tkiva&nbsp; u&nbsp; zrelom plodu i karakteristike receptakuluma predstavljaju karaktere koji su dijagnostički za rodove. Karakteri koji razdvajaju analizirane taksone&nbsp; jesu:&nbsp; karakteristike&nbsp; papusa&nbsp; kod&nbsp; vrsta&nbsp; roda&nbsp;<em> Pulicaria; </em>prstenasato&nbsp; suženje&nbsp; između&nbsp; baze&nbsp; papusa&nbsp; i&nbsp; cipsele,&nbsp; organizacija<br />sklerenhimskog&nbsp; tkiva&nbsp; ploda&nbsp; i&nbsp; tip&nbsp; trihoma&nbsp; na&nbsp; povr&scaron;ini receptakuluma&nbsp; kod&nbsp; vrsta&nbsp; roda<em>&nbsp; Dittrichia</em>;&nbsp; sekretorni&nbsp; kanali&nbsp; u perikarpu&nbsp; ploda,&nbsp; odsustvo&nbsp; trihoma&nbsp; i&nbsp; slabo&nbsp; izražena&nbsp; rebra&nbsp; na povr&scaron;ini&nbsp; receptakuluma&nbsp; kod&nbsp; vrste<em>&nbsp; L.&nbsp; crithmoides</em>.&nbsp; Plod&nbsp; vrste&nbsp;<em> I.</em><br /><em>helenium</em>&nbsp; razlikuje&nbsp; se&nbsp; od&nbsp; plodova&nbsp; svih&nbsp; preostalih&nbsp; analiziranih vrsta na osnovu njegovih dimenzija, većeg broja sklerenhimskih vrpci koje nisu istaknute u rebra, kao i na osnovu slabo izraženih rebara&nbsp; na&nbsp; receptakulumu,&nbsp; između&nbsp; karpopodijalnih&nbsp; otisaka,&nbsp; na kojima&nbsp; se&nbsp; nalaze&nbsp; žlezdane&nbsp; i&nbsp; nežlezdane&nbsp; trihome.&nbsp; Rezultati&nbsp; su<br />ukazali na&nbsp; to da se&nbsp; vrsta&nbsp; I. helenium&nbsp; izdvaja kao zaseban takson na osnovu anatomskih i mikromorfolo&scaron;kih karakteristika ploda i receptakuluma&nbsp; sa&nbsp; jednako&nbsp; malim&nbsp; stepenom&nbsp; statističke&nbsp; podr&scaron;ke koji&nbsp; takođe&nbsp; podržava&nbsp; izdvajanje&nbsp; rodova&nbsp; <em>Dittrichia,&nbsp; Limbarda&nbsp; i Pulicaria</em>&nbsp; u&nbsp; zasebne&nbsp; rodove.&nbsp; Stoga,&nbsp; na&scaron;i&nbsp; rezultati&nbsp; podržavaju tvrdnju&nbsp; da&nbsp; vrsta&nbsp; I.&nbsp; helenium&nbsp; treba&nbsp; biti&nbsp; izdvojena&nbsp; od&nbsp; preostalih<br />vrsta&nbsp; roda&nbsp; Inula&nbsp; u&nbsp; zaseban&nbsp; rod,&nbsp; pri&nbsp; čemu&nbsp; je&nbsp; njena&nbsp; autentičnost evidentna na nivou&nbsp; anatomije i mikromorfologije&nbsp; vegetativnih i reproduktivnih organa.<br />Rezultati&nbsp; hemijske&nbsp; analize&nbsp; etarskog&nbsp; ulja&nbsp; pružili&nbsp; su&nbsp; do&nbsp; sada nepoznate&nbsp; podatke&nbsp; o&nbsp; kvalitativnom&nbsp; i&nbsp; kvantitativnom&nbsp; sastavu etarskog ulja podzemnih organa analiziranih taksona i ukazali na mogućnost njihove primene u taksonomiji ispitivanog tribusa&nbsp; na nivou&nbsp; identifikacije&nbsp; pojedinačnih&nbsp; taksona.&nbsp; Takođe,&nbsp; dobijeni<br />rezultati&nbsp; ukazali&nbsp; su&nbsp; na&nbsp; nove&nbsp; potencijalno&nbsp; lekovite&nbsp; vrste&nbsp; koje predstavljaju&nbsp; potencijalni&nbsp; resurs&nbsp; za&nbsp; farmaceutsku&nbsp; industriju. Rezultati&nbsp; analize&nbsp; osnovnih&nbsp; komponenti&nbsp; kao&nbsp; i&nbsp; rezultati korespodentne&nbsp; analize&nbsp; ukazali&nbsp; su&nbsp; na&nbsp; odsustvo&nbsp; tendencije grupisanja&nbsp; taksona&nbsp; koji&nbsp; pripadaju&nbsp; istom&nbsp; rodu&nbsp; prema&nbsp; trenutno<br />važećoj&nbsp; klasifikaciji.&nbsp; Zbog&nbsp; velike&nbsp; varijabilnosti&nbsp; uslovljene faktorima&nbsp; spolja&scaron;nje&nbsp; sredine&nbsp; sastav&nbsp; etarskih&nbsp; ulja&nbsp; ne&nbsp; može&nbsp; se koristiti&nbsp; samostalno&nbsp; kao&nbsp; pouzdan&nbsp; hemotaksonomski&nbsp; marker. Ipak,&nbsp; rezultati&nbsp; o&nbsp; sastavu&nbsp; etarskih&nbsp; ulja&nbsp; značajni&nbsp; su&nbsp; u&nbsp; smislu njihove&nbsp; primene&nbsp; u&nbsp; medicinske&nbsp; i&nbsp; kozmetičke&nbsp; svrhe,&nbsp; u&nbsp; industriji<br />gde se etarska ulja koriste kao sirovina, te su dobijeni rezultati značajni&nbsp; i&nbsp; u&nbsp; kontekstu&nbsp; evaluacije&nbsp; kvaliteta&nbsp; etarskih&nbsp; ulja ispitivanih vrsta.<br />Sprovedeno&nbsp; istraživanje&nbsp; daje&nbsp; značajan&nbsp; doprinos&nbsp; u&nbsp; pogledu tačnije&nbsp; klasifikacije&nbsp; unutar&nbsp; tribusa&nbsp; koja&nbsp; se&nbsp; danas&nbsp; bazira uglavnom&nbsp; na&nbsp; morfolo&scaron;kim&nbsp; karakteristikama.&nbsp; Detaljno&nbsp; opisani morfo-anatomski&nbsp; i&nbsp; mikromorfolo&scaron;ki&nbsp; karakteri&nbsp; analiziranih<br />vegetativnih&nbsp; i&nbsp; reproduktivnih&nbsp; organa&nbsp; doprinose&nbsp; boljem poznavanju&nbsp; karakteristika&nbsp; vrsta&nbsp; analiziranih&nbsp; rodova&nbsp; i predstavljaju&nbsp; dopunu&nbsp; postojećim&nbsp; podacima&nbsp; o&nbsp; analiziranimtaksonima.&nbsp; Takođe,&nbsp; na&nbsp; bazi&nbsp; morfo-anatomskih&nbsp; karakteristika ploda i receptakuluma izrađen je dihotomi ključ za determinaciju analiziranih rodova i vrsta. Kori&scaron;ćenjem metoda diskriminantne i korespodentne analize izdvojeni su i karakteri na osnovu kojih je moguće&nbsp; izvr&scaron;iti&nbsp; diskriminaciju&nbsp; određenih&nbsp; grupa&nbsp; u&nbsp; okviru<br />analiziranih taksona.</p> / <p>A&nbsp; comparative&nbsp; analysis&nbsp; of&nbsp; morpho-anatomical&nbsp; and micromorphological characteristics of leaf, stem, rhizome, root, fruit and receptacle, as well as biochemical analysis of essential oils of underground organs, was conducted over 16 species of 4 genera,&nbsp;<em> Inula&nbsp; (I.&nbsp; aschersoniana,&nbsp; I.&nbsp; bifrons,&nbsp; I.&nbsp; britannica,&nbsp; I.conyza, I. ensifolia, I. germanica, I. helenium, I. oculus-christi, I. saslicina ssp. salicina, I. salicina ssp. aspera, I. spiraeifolia), Dittrichia&nbsp; (D.&nbsp; graveolens,&nbsp; D.&nbsp; viscosa),&nbsp; Limbarda&nbsp; (L. crithmoides)&nbsp; and&nbsp; Pulicaria&nbsp; (P.&nbsp; dysenterica,&nbsp; P.&nbsp; vulgaris)</em>&nbsp; tribe Inuleae,&nbsp; Compositae.&nbsp; In&nbsp; addition&nbsp; to&nbsp; the&nbsp; fact&nbsp; that&nbsp; the&nbsp; species from&nbsp; different&nbsp; genera&nbsp; analysed&nbsp; in&nbsp; this&nbsp; paper&nbsp; were&nbsp; previously placed&nbsp; in different sections within&nbsp; <em>Inula,&nbsp; I. helenium&nbsp; L</em>., as the type&nbsp; species&nbsp; of&nbsp; the&nbsp; genus&nbsp;<em> Inula,</em>&nbsp; has&nbsp; also&nbsp; been&nbsp; placed&nbsp; in&nbsp; a separate section&nbsp; Corvisartia.&nbsp; According to the latest molecular phylogenetic&nbsp; analyses,&nbsp; based&nbsp; on&nbsp; plastid&nbsp; and&nbsp; nuclear&nbsp; DNA sequence data, as well as morphological and karyological data, it&nbsp; is&nbsp; suggested&nbsp; that&nbsp; section&nbsp; Corvisartia&nbsp; should&nbsp; be&nbsp; segregated from&nbsp; the&nbsp; remaining&nbsp; Inula&nbsp; species&nbsp; as&nbsp; a&nbsp; separate&nbsp; genus,&nbsp; thus achieving&nbsp; a&nbsp; greater&nbsp; morphological&nbsp; homogeneity&nbsp; of&nbsp; the&nbsp; genus <em>Inula.&nbsp;</em> The objectives of dissertation are defined in accordance with&nbsp; the&nbsp; fact&nbsp; that&nbsp; the&nbsp; data&nbsp; on&nbsp; the&nbsp; morpho-anatomical, micromorphological and biochemical structure of the vegetative and reproductive organs of species of examined tribe are sparse and&nbsp; incomplete,&nbsp; while&nbsp; the&nbsp; analyses&nbsp; were&nbsp; carried&nbsp; out&nbsp; on&nbsp; few species&nbsp; of&nbsp; <em>Inula,&nbsp; Pulicaria,&nbsp; Dittrichia</em>&nbsp; and<em>&nbsp; Limbarda</em>,&nbsp; but&nbsp; less emphasis&nbsp; was&nbsp; placed&nbsp; on&nbsp; comparative&nbsp; purpose.&nbsp; The&nbsp; research involved&nbsp; a&nbsp; large&nbsp; number&nbsp; of&nbsp; qualitative&nbsp; and&nbsp; quantitative characters&nbsp; that&nbsp; were&nbsp; analysed&nbsp; and&nbsp; measured&nbsp; using&nbsp; a stereomicroscope,&nbsp; light,&nbsp; fluorescent&nbsp; and&nbsp; scanning&nbsp; electron microscope.&nbsp; The&nbsp; obtained&nbsp; data&nbsp; were&nbsp; statistically&nbsp; processed&nbsp; by various numerical analysis methods&nbsp; usually&nbsp; used in taxonomic research. These types&nbsp; of analyses for the species of the generaInula<em>,&nbsp; Dittrichia,&nbsp; Limbard&nbsp;</em> and&nbsp;<em> Pulicaria&nbsp;</em> have&nbsp; not&nbsp; been recorded in the literature so far. The&nbsp; conducted&nbsp; analysis&nbsp; showed&nbsp; that&nbsp; vegetative&nbsp; organs&nbsp; do&nbsp; not provide&nbsp; enough&nbsp; information&nbsp; on&nbsp; the&nbsp; basis&nbsp; of&nbsp; which&nbsp; certain groups&nbsp; of&nbsp; species&nbsp; could&nbsp; be&nbsp; singled&nbsp; out&nbsp; with&nbsp; statistically significant&nbsp; suport,&nbsp; neither&nbsp; the&nbsp; genera&nbsp; according&nbsp; to&nbsp; the&nbsp; actual classification.&nbsp; The&nbsp; yielded&nbsp; results&nbsp; indicated&nbsp; that&nbsp; the&nbsp; species from&nbsp; different&nbsp; genera&nbsp; were&nbsp; clearly&nbsp; delimited&nbsp; on&nbsp; the&nbsp; basis&nbsp; of their&nbsp; carpological&nbsp; and&nbsp; receptacular&nbsp; micromorphological&nbsp; and anatomical characteristics. Receptacle features and organisation of&nbsp; sclerenchyma&nbsp; tissue&nbsp; in&nbsp; a&nbsp; fruit&nbsp; are&nbsp; traits&nbsp; that&nbsp; tend&nbsp; to&nbsp; be diagnostic for genera. The features that separate analysed taxa&nbsp; are:&nbsp; two&nbsp; rows&nbsp; of&nbsp; pappus&nbsp; hairs&nbsp; in&nbsp;<em> Pulicaria&nbsp; </em>species;&nbsp; the&nbsp; ring constriction&nbsp; of&nbsp; the&nbsp; fruit&nbsp; over&nbsp; which&nbsp; it&nbsp; passes&nbsp; into&nbsp; pappus,&nbsp; the sclerenchymatous tissue in fruit in form&nbsp; of continuous ring, the type&nbsp; of&nbsp; trichomes&nbsp; on&nbsp; the&nbsp; receptacular&nbsp; surface&nbsp; in&nbsp;<em> Dittrichia </em>species;&nbsp; the&nbsp; fruit&nbsp; with&nbsp; secretory&nbsp; ducts,&nbsp; and&nbsp; the&nbsp; receptacular surface&nbsp; with&nbsp; weakly&nbsp; pronounced&nbsp; ridges&nbsp; without&nbsp; glands&nbsp; in&nbsp; L. crithmoides.&nbsp; I. helenium&nbsp; fruits differ from those of all the other species&nbsp; by&nbsp; their&nbsp; size,&nbsp; greater&nbsp; number&nbsp; of&nbsp; inconspicuous sclerenchymatic&nbsp; bundles,&nbsp; and&nbsp; poorly&nbsp; expressed&nbsp; ridges&nbsp; among carpopodial prints&nbsp; with glandular and non-glandular trichomes distributed along them. Results revealed that I. helenium&nbsp; stands out&nbsp; as&nbsp; a&nbsp; separate&nbsp; taxon,&nbsp; based&nbsp;&nbsp; on&nbsp; the&nbsp; anatomical&nbsp; and micromorphological&nbsp; characteristics&nbsp; of&nbsp; its&nbsp; fruit&nbsp; and&nbsp; receptacle, with equally small percentage of statistical support, which also supports the recognition of<em> Pulicaria,&nbsp;</em> <em>Dittrichia</em> and<em> Limabarda</em> as&nbsp; separate&nbsp; genera.&nbsp; Obtained&nbsp;&nbsp; results&nbsp; support&nbsp; the&nbsp; claim&nbsp; that&nbsp; I. helenium should be separated from the Inula genus, whereby its authenticity&nbsp; is&nbsp; evident&nbsp; at&nbsp; the&nbsp; level&nbsp; of&nbsp; anatomy&nbsp; and&nbsp; micromorphology of vegetative and reproductive organs. The chemical analysis of the essential oil has provided new&nbsp; data on the qualitative and quantitative composition of the essential oil of the&nbsp; underground organs of the analysed taxa&nbsp; and pointed out&nbsp; the&nbsp; possibility&nbsp; of&nbsp; their&nbsp; application&nbsp; in&nbsp; taxonomy.&nbsp; Also,&nbsp; the obtained results indicated potentially&nbsp; new medicinal plants that represent&nbsp; a&nbsp; potential&nbsp; resource for the&nbsp; pharmaceutical&nbsp; industry. Principal&nbsp; component&nbsp; analysis&nbsp; as&nbsp; well&nbsp; as&nbsp; the&nbsp; results&nbsp; of&nbsp; the Correspondent&nbsp; analysis&nbsp; indicated&nbsp; the&nbsp; absence&nbsp; of&nbsp; the&nbsp; grouping tendency of taxa belonging to the same genus,&nbsp; according to the actual classification. Due to the high variability conditioned by the&nbsp; environmental&nbsp; factors&nbsp; the&nbsp; composition&nbsp; of&nbsp; essential&nbsp; oils cannot&nbsp; be&nbsp; used&nbsp; independently&nbsp; as&nbsp; a&nbsp; reliable&nbsp; chemotaxonomic&nbsp; marker.&nbsp; However,&nbsp; the&nbsp; results&nbsp; of&nbsp; the&nbsp; composition&nbsp; of&nbsp; essential oils are significant in terms of their application for medical and cosmetic&nbsp; purposes, in the industry&nbsp; where&nbsp; oils&nbsp; are&nbsp; used&nbsp; as raw materials,&nbsp; and&nbsp; the&nbsp; results&nbsp; obtained&nbsp; are&nbsp; also&nbsp; significant&nbsp; in&nbsp; the context&nbsp; of&nbsp; evaluation&nbsp; of&nbsp; the&nbsp; quality&nbsp; of&nbsp; investigated&nbsp; essential oils.The&nbsp; conducted&nbsp; research&nbsp; makes&nbsp; a&nbsp; significant&nbsp; contribution&nbsp; to more precise classification within the&nbsp; tribe, which is now basedmainly&nbsp; on&nbsp; morphological&nbsp; characteristics.&nbsp; The&nbsp; detailed description&nbsp; of&nbsp; morpho-anatomical&nbsp; and&nbsp;&nbsp; micromorphological characters&nbsp; of&nbsp; the&nbsp; analysed&nbsp; vegetative&nbsp; and&nbsp; reproductive&nbsp; organs contributes&nbsp; to&nbsp; better&nbsp; knowledge&nbsp; of&nbsp; the&nbsp; characteristics&nbsp; of&nbsp; the analysed&nbsp; species&nbsp; and&nbsp; complements&nbsp; the&nbsp; existing&nbsp; data&nbsp; on&nbsp; them. Also, on the basis of morpho-anatomical characteristics of the fruit and the receptacle a dichotomous key&nbsp; for the determination of the analysed genera and species&nbsp; was developed. According to&nbsp; the&nbsp; Discriminant&nbsp; and&nbsp; Correspodent&nbsp; analysis&nbsp; the characteristics&nbsp; on&nbsp; which&nbsp; is&nbsp; possible&nbsp; to&nbsp; discriminate&nbsp; certain groups within the analysed genera are also distinguished.</p>

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1. https://www.cris.uns.ac.rs/DownloadFileServlet/Disertacija152810962145537.pdf?controlNumber=(BISIS)107459&fileName=152810962145537.pdf&id=11485&source=NDLTD&language=en
2. https://www.cris.uns.ac.rs/record.jsf?recordId=107459&source=NDLTD&language=en

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Identifer	oai:union.ndltd.org:uns.ac.rs/oai:CRISUNS:(BISIS)107459
Date	27 September 2017
Creators	Karanović Dunja
Contributors	Zorić Lana, Luković Jadranka, Anačkov Goran, Orčić Dejan, Zlatković Bojan
Publisher	Univerzitet u Novom Sadu, Prirodno-matematički fakultet u Novom Sadu, University of Novi Sad, Faculty of Sciences at Novi Sad
Source Sets	University of Novi Sad
Language	Serbian
Detected Language	Unknown
Type	PhD thesis