<p><em>Prospero autumnale </em> kompleks je taksonomski najintrigantniji član roda <em>Prospero,</em> sa centrom rasprostranjenja u Mediteranu. Kompleks se rasprostire i na obalama Atlantskog okeana u Francuskoj, sve do južnih delova Velike Britanije, zatim na Balkanskom poluostrvu, u Panonskoj niziji, sve do Krima, Kavkaza i delova Irana na istoku. Areal kompleksa se preklapa sa arealima ostale dve vrste roda – u<br />zapadnom Mediteranu sa <em>P. obtusifolium</em>, a u istočnim delovima Mediterana sa <em> P. hanburyi. </em>Za razliku od <em>P. obtusifolium i P. hanburyi, P. autumnale</em> kompleks se odlikuje visokom kariološkom varijabilnošću.Razlikuju se četiri osnovna broja hromozoma x = 5, 6, 7 i četiri različita tipa genoma (A, B<sup> 5 </sup>, B <sup>6</sup> i B<sup> 7</sup> ) i diploidna citotipa (AA, B <sup>5 </sup>B<sup> 5 </sup>, B <sup>6 </sup>B<sup> 6 </sup>i B <sup>7 </sup>B <sup>7</sup> ) koja se razlikuju u odnosu na osnovni broj hromozoma, veličinu i morfologiju hromozoma. Genom B <sup>5</sup> ima x = 5, B<sup> 6</sup> x = 6, a genomi A i B<sup> 7 </sup>x = 7. Najrasprostranjeniji genom je B <sup>7</sup> , koji je zabeležen u celom arealu kompleksa, dok je A zastupljen u zapadnom Mediteranu, B<sup> 5</sup> u Libiji, a B <sup>6 </sup>je endemičan za Krit. U okviru B <sup>7</sup> genoma se, dalje, razlikuju dve linije nastale kao posledica duplikacija 5S rDNK lokusa, pri čemu linija I ima jedan lokus, a kod linije II taj lokus je duplikovan. Osim diploidnih citotipova, poznato je i nekoliko ploidnih nivoa, od kojih su najučestaliji tetraploidi (2n=4x=28) i heksaploidi (2n=6x=42). Za <em>P. autumnale </em> kompleks je karakteristična varijabilnost u veličini genoma između različitih citotipova i unutar pojedinih citotipova. Genomsko restrukturiranje, koje je imalo najveću ulogu u evoluciji i diverzifikaciji kompleksa, nije imalo velikog uticaja na morfološku varijabilnost, te je iz tog razloga kompleks morfološki skoro uniforman. Ipak karakteri kao što su: boja tunike lukovice, visina biljke, oblik i dimenzije lista, broj cvetova u cvasti i boja cvetova pokazuju izvesnu varijabilnost. Taksonomske nejasnoće nastaju zbog opisanih desetak vrsta između kojih ne postoje jasne morfološke razlike i kod kojih je prisutno preklapanje vrednosti karaktera.Kako je do sada kompleks najviše bio predmet karioloških istraživanja, pri čemu su izostale detaljne analize morfološke i/ili anatomske varijabilnosti, uzorkovan je biljni materijal sa 37 lokaliteta na području Panonske nizije i Balkanskog poluostrva sa ciljem utvrđivanja morfološke, anatomske, kao i kariološke varijabilnosti. Određivan je broj hromozoma, nivoi ploidije i veličina genoma. Primenom univarijantne statističke metode, ispitana je varijabilnost morfoloških i anatomskih karaktera, a upotrebom multivarijantne statističke metode su testirane razlike između unapred definisanih grupa, koje su se odnosile na populacije i ploidne nivoe. Korespodentnom analizom su analizirani kvalitativni morfološki i anatomski karakteri. Ukupno je analizirano 65 karaktera (33 morfološka i 32 ana tomska). Rezultati kariološke analize se potvrdili varijabilnost kompleksa na istraživanom području. Detektovana su tri ploidna nivoa (diploidi, tetraploidi i heksaploidi), a potvrđena je i varijabilnost u veličini genoma. Poređenjem monoploidne veličine genoma između tri ploidna nivoa je detektovano smanjivanje veličine genoma sa povećanjem nivoa ploidije. U odnosu na koeficijent varijabilnosti, konstatovano je da se većina morfoloških karaktera nalazi u zoni umerene varijabilnosti (22), a samo pet karaktera je visokovarijabilno. Anatomski karakteri su raspoređeni u četiri kategorije (niskovarijabilni, umerenovarijabilni, visokovarijabilni i veoma visokovarijabilni), pri čemu većina karaktera (20) pripada umerenovarijabilnoj kategoriji. Četiri kvalitativna morfološka (oblik lukovice, boja tunike, boja cveta i oblik plodnika) i sedam kvalitativnih anatomskih karaktera (oblik poprečnog preseka lista, oblik ćelija epidermisa na licu i naličju, oblik ćelija palisadnog tkiva na licu i naličju, prisustvo papila i kristala u ćelijama parenhima) su pokazala varijabilnost. U korespodentnoj analizi, zasnovanoj na kvalitativnim morfološkim karakterima, izdvojile su se tri grupe. Za prvu grupu su bili zajedničke širokojajaste i uzanojajaste lukovice sa ružičastom tunikom, jedinkama u drugoj grupi su bili svojstveni jajasti i uzanojajasti plodnici i lukovice sa braon tunikom, dok su za treću grupu bili karakteristični širokojajasti plodnici, loptaste i pljosnate lukovice. U odnosu na kvalitativne anatomske karaktere najviš e se izdvojila jedna grupa za koju su bile karakteristične gusto raspoređene papile i prisustvo četvorougaonih i okruglastih epidermalnih ćelija na abaksijalnoj strani lista. Rezultati multivarijantnih analiza, zasnovanih na populacijama kao unapred definisanim grupama i kvantitivnim karakterima, ukazali su na složenu varijabilnost uzorka i izostanak jasne separacije grupa. Naznake razdvajanja grupa su bile uočljive u analizi morfo-anatomske matrice. Izdvojili su se sledeći karakteri: visina stabla, dužina cvasti, broj cvetova, površina poprečnog preseka lista, ukupna površina palisadnog tkiva i ukupna površina sunđerastog tkiva. Najjasnija razdvajanja su uočena u analizama gde su ploidni nivoi predstavljali a priori definisane grupe, naročito u kombinaciji morfoloških i anatomskih karaktera. Karakteri sa diskriminacionim potencijalom između jedinki koje pripadaju različitim ploidnim nivoima jesu: broj<br />cvetova, širina listića perigona unutrašnjeg kruga i filamenta spoljašnjeg kruga, površina poprečnog preseka lista, ukupna površina palisadnog tkiva, površina ćelija palisadnog tkiva, visina i širina ćelija palisadnog tkiva i udeo epidermisa. Od sva tri ploidna nivoa, najviše izdvojene su bile diploidne jedinke, koje se od tetraploida i heksaploida mogu razlikovati na osnovu: dužina filamenata spoljašnjeg i unutrašnjeg kruga cveta, površine ćelija palisadnog tkiva, visine i širine ćelija palisadnog tkiva, kao i ukupne površine palisadnog tkiva. Tetraploidi i heksaplodi se najviše međusobno razlikuju na osnovu karaktera u regionu cveta: prečnik otvorenog perigona, dužine listića perigona spoljašnjeg kruga cveta i širine filamenta spoljašnjeg kruga cveta. Uočene razlike između tri analizirana ploidna nivoa (diploidi, tetraploidi i heksaploidi), u taksonomskom smislu se mogu tumačiti pre kao jedna od infraspecijskih kategorija, a ne kao kategorija koja bi odgovarala vrsti.</p> / <p>Prospero autumnale complex is the most taxonomically intricate member of the genus Prospero with the centre of distribution on the Mediterranean Basin, westwards to the Atlantic coast in France as far north as Great Britain, northwards to the Pannonian Basin and Crimea, and eastwards to Iran. The distribution area of the complex overlaps with distribution areas of the other two species of the genus<br />Prospero-P. obtusifolium is present in the western parts of the Mediterranean, while P. hanburyi is distributed in the eastern Mediterranean. Unlike P. obtusifolium and P. hanburyi, P. autumnale complexis characterized by extraordinary karyological cariation. It encompasses four distinct genomes (A, B 5 , B 6 , B 7 ) and four distinct diploid cytotypes (AA, B 5 B 5 , B 6 B 6 , B 7 B 7 ), each with a unique combination of<br />basic chromosome number (x = 5, 6, 7). An additional difference is related to chromosome size and morphology. The B 5 genome has x = 5, B 6 x = 6, and A and B 7x = 7 basic chromosome numbers. The B 7 genome is present in the whole distribution area of P. autumnale complex, while genome A is distributed in the western Mediterranean, B 5 is restricted to Libya, and B 6 is endemic to Crete. Within diploids and tetraploids of B 7 genome, two lineages occur as a consequence of duplication of the 5S rDNA locus. Type I (lineage) has a single locus, while in type II (lineage) 5S rDNA locus is duplicated. Besides diploid cytotypes, the polyploids cytotypes are also known. The most common polyploids are tetra- (2n=4x=28) and hexaploids (2n=6x=42). Variability in the genome size within, as well as between cytotypes, is also characterized for the complex. Genomic restructuring, which played the b iggest role in evolution and diversification of the P. autumnale complex, did not have a major impact on morphological variability. Only slight variation has been detected in plant, tepal and filament size, flower and seed color, shape and size of leaves a nd color of bulbs. Taxonomic ambiguities are<br />additionally caused by description of the new ten species of the complex. morphological differences between those species are unclear, with overlapping values of the most morphological traits. Until now, P. autumnale complex was mostly karyologicaly investigated, without detail analyses of morphological and/or anatomical variability. The aim of the present study was to investigate karyological, morphological and anatomical variability on individuals collected on 37 localities across the Pannonian Basin and Balkan Peninsula. Basic chromosome number, ploidy levels, as well as genome size were determined. The variability of morphological and anatomical characters was examined using univariate statistical methods. Differences between predefined groups (populations and ploidy levels) were investigated using multivariate statistics with the attempts to identify morphological and anatomical characters with discriminatory potential. For analysing qualitative morphological and anatomical characters, correspondence analysis was conducted. In total 65 traits were analyzed (33 morphological and 32 anatomical). The results of karyological analysis confirmed the high variation of the P. autumnale complex in the study area. Three ploidy levels (diploids, tetraploids and hexaploids) were detected, and the genome size variation was confirmed. Genome downsizing was observed by comparing monoploid genome sizes between three ploidy levels. According to the oefficient ofc variation, most morphological characters show moderate variation (22). Only five traits showed high variation. Anatomical characters were classified into four categories according to the coefficient of variation (with low, moderate, high and very high variation), but the most traits (20) showed moderate<br />variation. Variation in qualitative morphological (bulb shape and color, flower color and ovary shape) and seven qualitative anatomical characters (leaf cross-sectional area shape, shape of the adaxial and abaxial epidermal cells, shape of the adaxial and abaxial palisade cells, presence of the papillae and crystals in the parenchyma cells) were recorded. Three groups of populations, as a result of the correspondence analysis, based on qualitative morphological characters, were formed. Populations of the first group showed common characteristics such as broadly and narrowly ovate shaped bulbs with pink outer tunic. The second group was characterized by bulbs with brown outer tunic, ovate and narrowly ovate ovary, while the third group had broadly ovate ovaries, globose and flattened bulbs. In relation to qualitative anatomical characters, only one group was the most distinguished and was characterized by densely distributed papillae, squared and rounded shape of abaxial epidermal cells. The results of multivariate analyses of quantitative traits, based on populations as presumptive groups, revealed similar patterns without structuring and with no specific groupings. The tendency toward a separation of the populations was noticeable in the analysis based on combined morpho-anatomical characters. The highest correlation with the canonical axes showed: stem height, inflorescence length,number of flowers, leaf cross-sectional area, palisade and spongy tissue area. The clearest separationsbetween groups were observed with ploidy levels as presumptive groups. Morpho-anatomical traits with discriminatory potential among ploidy levels were: number of flowers, inner tepal width, outer filament width, leaf cross-sectional area, palisade tissue area, the cross-sectional area of palisade cells, height and width of palisade cells and epidermis percentage. The most distinct group among three ploidy levels were diploids and could be distinguished from tetra- and hexaploids by outer and inner filament width, the cross-sectional area of palisade cells, the width of palisade cells, the height of palisade cells and palisade tissue area. Tetra- and hexaploids differed mostly in floral characters: open flower diameter, outer tepal length and outer filament width. For taxonomic purposes, the level of overlap indicates that the ploidy levels could be regarded as showing intraspecific variation.</p>
Identifer | oai:union.ndltd.org:uns.ac.rs/oai:CRISUNS:(BISIS)110998 |
Date | 18 October 2019 |
Creators | Vestek Ana |
Contributors | Anačkov Goran, Luković Jadranka, Zlatković Bojan, Šinžar-Sekulić Jasmina, Lazarević Maja |
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 |
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