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Dendrobiology 59.Vp 2008, vol. 59, 13–22 Heike Liesebach, Zoltán Sinkó A contribution to the systematics of the genus Tilia with respect to some hybrids by RAPD analysis Received: 02 January 2008, Accepted: 28 March 2008 Abstract: The putative hybridcharacter of two Tilia varieties selectedas avenue trees (‘Szent Istvan’ and‘K3’) shouldbe clarifiedfor further breedingactivities. Their ancestor species shouldbe identifiedbya genetic comparison with reference material (Tilia cordata, T. platyphyllos, T. dasystyla, T. × euchlora, T. × europaea and others). RAPD marker data were evaluated by UPGMA cluster and neighbour-joining analysis. The genetic comparison of the two selectedclones with the collection of reference material offers insights into some sys- tematic relationships within the genus Tilia. It was supplemented by a critical discussion of methods of RAPD data evaluation for taxonomic purposes. Additional key words: Tilia L.; linden; taxonomy; hybrids; DNA; UPGMA cluster analysis; neighbour-join- ing; resampling Addresses: H. Liesebach, Institute of Forest Genetics, Johann Heinrich von Thünen-Institute (vTI), Federal Research Institute for Rural Areas, Forestry and Fisheries, Eberswalder Chaussee 3A, 15377 Waldsievers- dorf, Germany, e-mail: [email protected] Z. Sinkó, Nursery Prenor, Béke tér 1, 9707 Szombathely-Herény, Hungary Introduction plantings were undertaken more for ornamental rea- sons. Since the 1960s, T. cordata Miller and T. Trees andshrubs of the genus Tilia L. (Malvaceae) platyphyllos Scop. were increasingly replacedby T. were foundin the forests of the northern hemisphere tomentosa Moench and T. × euchlora K. Koch from in Europe, Asia andEastern North America andCen - southeast Europe andthe Caucasian region because tral America. They are growing in temperate, sub- of their better adequacy to urban conditions in Cen- tropical andtropical climates andoccur from moist to tral Europe. Actually the breeding efforts focus on dry regions(Muir 1984). better-adapted plant material with high drought and Representatives of these deciduous trees became salt resistance andmore tolerance to soil compaction increasingly important in municipal parks of large cit- for urban locations (Karnosky et al. 1982). ies andplay a central role as avenue trees (Sukopp and In Budapest, several trees from an avenue with Wurzel 2000). Growing damages of lime-trees were positive traits in stress resistance were selected observedin European cities because of the ubiquitous (Jámborné et al. 2001). They are about 30 years old stress factors as salt anddrought[Helsinki: Terho andoriginate from seedlingoffsprings after free polli - andHallaksela (2005), Warsaw: Chmielewski et al. nation. They are putative hybrids but their parents (1998), Budapest: Sinkó (unpublished)]. Former couldnot be reconstructed.Presumedancestors seem 14 Heike Liesebach, Zoltán Sinkó to originate from the group of European andCauca - reproducibility was accepted. For the application of sian lime-trees. One selectedvariety (‘S zent István’) RAPD markers no molecular information is needed develops morphological characteristics similar to T. for short, arbitrary primers. As a fingerprinting dasystyla Steven or T. × euchlora. The other variety methodthese markers are well establishedfor clonal (‘K3’) is more similar to T. platyphyllos. However, their andcultivar identificationof numerous plant species. morphological characteristics of twigs, leaves, flowers Representative examples amongst others are given andfruits didnotallow a doubtlesstaxonomic classifi - for poplar, melon and Citrus (Rajora andRahman cation to fulfil the standards for traded ornamental 2003; Tanaka et al. 2007; Rao et al. 2007). By applica- plants (Sinkó 2005). The morphological findings of tion of a sufficient number of primers andevaluable the selectedclones of interest shouldnow be com - bands, RAPDs reveal good estimations of genetic pletedby genetic datato get more information on their similarities and degree of relationship. taxonomic status for further breeding. This suggestion bases on the hypothesis that phylogenetic relation- Methods ships couldbe reflectedby suitable genetic markers andcomparedto adequatereference material. Several Plant material methods of data evaluation should be critically re- The putative hybridclone ‘Szent István’ (No. 1 in viewedto reconstruct the evolutionary relationships of Table 1) andthe clone ‘K3’ (No. 2), which is very the investigatedgroup of organisms. close to T. platyphyllos regarding morphological traits, Only some genetic studies in the genus Tilia (T. are selectedclones for urban locations with a prob - cordata and T. platyphyllos) were carriedout by lematic taxonomic status. A collection of plant mate- isozyme markers (Maurer andTabel 1995; Fromm rial from the genus Tilia was compiledto cover the pu - andHattemer 2003). Isozyme markers in the context tative relatives, which were assumedamong the Eu- of population genetic studies of outcrossing species ropean andCaucasian group of Tilia species basedon are known to possess a high level of within popula- morphological observations (T. dasystyla, T. dasystyla tion variation. An additional difficulty for codominant Steven subsp. caucasica (V. Engl.) Pigott and T. × nuclear markers is the polyploidisation in the phylo- euchlora). T. × euchlora is a putative hybridof T. cordata geny of these lindens (Fromm and Hattemer 2003). and T. dasystyla. However, sufficient population basedmaterial from A number of individuals of available species were the Caucasian species group couldnot be provided included to estimate within and among population andthus isozyme markers didnotcome into ques- variation at the species level as far as possible (T. tion. Nuclear microsatellite markers for Tilia species cordata and T. platyphyllos). The specimens No. 26 and are not available so far. Even if they existed, they No. 27 are labelledas T. dasystyla in the Botanical Gar- wouldnot be suitable for our studyfor the same rea - den Berlin-Dahlem, but they were identified as T. son like isozyme markers. cordata by morphological traits (Sinkó unpublished). High variation in chloroplast DNA haplotypes was Two other species with only 3 individuals were in- detected for European T. cordata populations (Fi- cluded into the experiment: T. tomentosa and T. neschi et al. 2003). However, chloroplasts were amurensis Rupr. mostly maternal inheritedin Angiosperms (Harris Furthermore some selectedclones were involved: andIngram 1991; Rajora andDancik 1992) andthere - T. cordata ‘Wega’, T. tomentosa ‘Szeleste’, T. × europaea fore did not reveal taxonomic relationships in case of L. ‘Pallida’, and one unknown Tilia specimen from the possible hybridisation between closely related spe- Späth Arboretum in Berlin. T. × europaea is possibly a cies. Secondary, it is known that several related spe- hybrid of T. cordata and T. platyphyllos. cies share their chloroplast genotypes within the ge- In addition, species far distant from the putative nus [i.e. Tilia (Brunner et al. 2001), Quercus L. relatives of the interesting clones were integratedas (Dumolin-Lapegue et al. 1997), Betula L. (Palme et al. outgroups to test the reliability of the method(Amer - 2004), Hedera L. (Grivet andPetit 2002)]. Thus, ica: T. americana L., Asia: T. henryana Szyszyll.). chloroplast markers are not appropriate for our sub- ject. RAPD analysis Markers widely distributed within the genome Twigs were harvestedin February andMarch 2004. wouldbe the methodof choice for visualisation of They were cultivatedin greenhouse up to the devel - phylogenetic relationships andto get information on opment of first young leaves. Leaves were frozen in the origin of putative hybrids, even if the number of liquidnitrogen andhomogenised.DNA was isolated reference accessions is low: species-independent with the help of DNeasy Plant Mini Kit (QIAGEN) ac- RAPD (Random Amplified Polymorphic DNA) and cording to manufactures instructions. The samples of AFLP markers (AmplifiedFragment Length Polymor - 20–50 mg leaf tissue yielded in respectively 100–250 phism). Due to the fact that AFLPs needan extensive ng DNA estimated by photometric assay. laboratory input, the risk of somewhat lower A contribution to the systematics of the genus Tilia with respect to some hybrids by RAPD analysis 15 A standard protocol was used to carry out the minutes at 95°C; 30 cycles with 30 sec 95°C denatur- RAPD analysis. A total of 36 arbitrary primers were ation, 30 sec 35°C annealing, 1 min 72°C elongation; usedfor the amplification (Table 2). Each PCR (Poly - final extension 7 min 72°C; hold 4°C. merase Chain Reaction) was performedin 25 µl reac - PCR products were separated in 1.8% agarose gel tion volume containing 100–250 pg template DNA, by electrophoresis andstainedwith ethidiumbro - dNTP mix (200 µM each nucleotide), 0.4 µM primer, mide. The gels were photographed using the Kodak reaction buffer (including MgCl2 to a final concentra- Edas 290 System. tion of 2.5 mM) and1 Unit Taq polymerase (Amer - A replication of DNA extraction was carriedout sham Pharmacia Biotech, NJ, USA). with tissues from buds and cambium in one case and The amplification reactions were carriedout in a replications of DNA amplification were performed thermocycler (Biometra GmbH, Göttingen, Germa- with 17 relevant primers anda subset of samples to ny) with the following program: Initial denaturation 5 confirm reproducibility. Table 1. Material collection of genus Tilia including selected clones No. Species and/or cultivar Abbreviation Location
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