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Comparison of Three Polygonatum Species from Poland Based on DNA Markers

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Comparison of Three Polygonatum Species from Poland Based on DNA Markers Ann. Bot. Fennici 43: 379–388 ISSN 0003-3847 Helsinki 24 October 2006 © Finnish Zoological and Botanical Publishing Board 2006 Comparison of three Polygonatum species from Poland based on DNA markers Monika Szczecińska1*, Jakub Sawicki1, Kornelia Polok2, Czesław Hołdyński & Roman Zieliński2 1) Department of Botany and Nature Protection, University of Warmia and Mazury, Plac Łódzki 1, PL-10-727 Olsztyn, Poland (*corresponding author’s e-mail: [email protected]) 2) Department of Genetics, University of Warmia and Mazury, Plac Łódzki 3, PL-10-967 Olsztyn, Poland Received 1 Sep. 2005, revised version received 12 Feb. 2006, accepted 23 Mar. 2006 Szczecińska, M., Sawicki, J., Polok, K., Hołdyński, C. & Zieliński, R. 2006: Comparison of three Polygonatum species from Poland based on DNA markers. — Ann. Bot. Fennici 43: 379–388. Four categories of DNA markers were used to determine genetic similarity of three spe- cies of Polygonatum, i.e. P. multiflorum, P. odoratum and P. verticillatum, occurring in Poland. Four populations per species, represented by ten plants per population, were collected in north-eastern Poland. In this study 111 RAPD and 35 semi-random ISJ markers were detected. Moreover the application of primers complementary to bacterial sequences IS6110 and katG gene enabled the detection of 9 and 34 markers, respec- tively. Each of the four marker categories gave species-specific bands. The highest conservativeness was observed for sequences of the katG gene. The degree of genetic similarity between P. multiflorum and P. odoratum was 0.57, and between these taxa and P. verticillatum 0.40 and 0.37, respectively. Most of the primers applied in the study ena- bled molecular species identification, and three of them turned out to be section-specific markers. The taxa showed no considerable intraspecific diversity, and the populations of P. verticillatum were almost identical. The results are consistent with the commonly accepted division of the genus Polygonatum into sections Polygonatum and Verticillata. Key words: DNA markers, genetic similarity, Polygonatum Introduction analyses of enzymes, concluded that the index of genetic similarity characterizing good biological In taxonomy, morphological markers constitute species should be lower than 0.6. Such species the main criteria of species distinction. By using have reproductively isolated genetic pools and molecular techniques we have the possibility to typically show the presence of many specific validate the taxonomic classification, because markers allowing their easy identification. the species-specific morphological differences However, “botanical” species do not always should correspond to certain differences on the turn out to be good “biological” species. Two molecular level. Hamrick and Godt (1990), in species of grass, Lolium perenne and L. mul- their summary of over 20 years of electrophoretic tiflorum may serve as an example since they 380 Szczecińska et al. • ANN. BOT. FENNICI Vol. 43 do not have enzymatic and DNA markers and been studied with DNA markers yet. This study have the genetic similarity of 0.96 (Polok 2005). also gives the possibility to determine, for the The enzymatic markers, despite numerous exam- first time, the levels of their intra- and inter-spe- ples of their application for taxonomic purposes, cific variation. seem to be inferior to DNA markers, which can be generated in any number and can represent different genomic sequences. For example, spe- Material and methods cies-specific enzymatic markers have not been found in Scots pine, Pinus sylvestris or dwarf Material pine, P. mugo. However, these pines have differ- ent DNA markers of both unique and RAPD-type Polygonatum belongs to the Convallariaceae and nonspecific sequences (Zieliński & Polok 2005). has a total of 57 species. Its major distribution A sibling species, enzymatically undistinguish- centers are in East Asia, mainly China and Japan, able from the typical form of the liverwort Pellia where 40 species are found (Tamura 1997). Apart endiviifolia, has been identified similarly (Polok from this area, Polygonatum occurs in the moder- et al. 2005b). ate climate zone of North America and Europe. The usefulness of different DNA sequences of Morphometric and cytological analyses per- nuclear and organellar genomes has been tested formed so far (Therman 1957, Nowakowska and in molecular studies for taxonomic purposes. Żeglicka 1972, Abramova 1975, Tamura 1993) The bacterial sequences present in the eukaryo- on Polygonatum enabled two sections to be dis- tic genome characterized by high amplification tinguished: Polygonatum and Verticillata. Section specificity have been also used recently. The Polygonatum is characterized by leaves arranged sequences complementary to the bacterial gene alternately on the stem, thick filaments, perfo- katG coding catalase-peroxidase enzyme and to rated pollen grains and chromosome number x the bacterial insertion sequence IS6110 belong = 9, 10 or 11. Their karyotypes contain mostly to this category. Employing these sequences ena- metacentric and submetacentric chromosomes. bled development of many markers identifying The section Verticillata is characterized each of the 16 analyzed crop cereals and grass by verticillate leaves, thin filaments, reticulate species (except for Lolium perenne and L. mul- pollen grains and chromosome number x = 14 or tiflorum), selected species of the genus Pinus, as 15. Karyotypes contain mostly long subtelocen- well as liverwort species (Chmiel & Polok 2005, tric and short metacentric chromosomes (Tamura Zieliński & Polok 2005). 1993). The purpose of this study was to test whether Additional information was provided by the two bacterial sequences described above Tamura (1997), who investigated the chloroplast could be used to identify selected species of DNA to estimate the phylogenetic relationships dicotyledon plants. The effectiveness of the between 23 species of Polygonatum. The phy- above markers in species identification was logenetic tree created on the basis of these data tested on three species of Polygonatum found in confirmed the earlier division of the genus into Poland: P. multiflorum, P. odoratum and P. verti- two sections, as well as the classification of par- cillatum. These species constitute a good model ticular taxa into these sections, based mostly on because they are easy to distinguish morphologi- cytological and morphological analyses. cally and their taxonomic status raises no doubts. In Poland Polygonatum is represented by The species belong to two sections, Polygo- three species, i.e. P. multiflorum and P. odoratum natum and Verticillata, therefore the bacterial of the section Polygonatum, and P. verticillatum sequences can be used in distinguishing both of the section Verticillata. Polygonatum multiflo- species and sections. The bacterial sequences rum and P. odoratum can be found in all parts of katG and IS6110 were assessed in the context of Poland, but in different habitats. The former is widely used markers RAPD and semi-specific reported mainly from dry-ground forests, and the markers ISJ (Intron Splice Junction). latter from pine and mixed coniferous forests, Species of the genus Polygonatum have not and oak forests (Matuszkiewicz 2001). Polygo- ANN. BOT. FENNICI Vol. 43 • Comparison of three Polygonatum species based on DNA markers 381 natum verticillatum, considered a montane spe- RAPD markers cies (Zając 1996), has a disjunctive distribution in Poland. It can be found in both mountain areas RAPD markers are universally applied to deter- and lowlands of northern Poland. In the moun- mine genetic diversity within plant and animal tains it usually occurs in forest communities populations (Williams 1990, Stenoien 1999, of the class Vaccinio–Piceetea and grass com- Polok et al. 2005b). In contrast to specific mark- munities of the alliance Calamagrostion. In the ers, RAPD markers are randomly distributed lowlands it is most common in shaded forests of in the genome, thus permitting an objective the alliances Carpinion and Fagion. assessment of genetic polymorphism of a given The studied material comprised four popula- species, without the need to know the func- tion samples of each species, P. odoratum, P. tions and structure of the amplified sequence, multiflorum and P. verticillatum, collected in which significantly reduces the costs of analysis. northeastern Poland. The localities are shown in The sequences of RAPD primers used for DNA Table 1. Each sample consisted of ten plants ran- amplification in this study are given in Table 2. domly collected from the population. The leaves The PCR reaction was conducted in a volume from each plant were cleaned and then stored at of 20 µl containing 1 µl PCR buffer (400 mM –20 °C. (NH4)2SO4 and 1 M Tris-HCl, pH 9 at 25 °C), 2 mM MgCl2, Enhancer containing betaine, 200 µM each dATP, dGTP, dCTP, dTTP, 0.3 µM DNA extraction primer, 1 unit of Tfl polymerase (Epicentre Tech- nology) and 80 ng of template DNA. The reac- In total 1 g of leaves was taken from ten indi- tion was carried out at 94 °C for 3 min. followed viduals of each population. by 45 cycles at 94 °C for 1 min., 36 °C for 1 DNA was isolated by the modified CTAB min., and 72 °C for 2.5 min., with a final exten- procedure. The liquid nitrogen-ground leaves sion step of 72 °C for 5 min. were thoroughly mixed with 3 ml preheated CTAB isolation buffer (2% CTAB, 100 mM Tris- HCl, pH 8.0, 20 mM EDTA, 1.4 M NaCl and ISJ markers 2% β-mercaptoethanol) and incubated at 55 °C for 1 h. After three chloroform extractions, the Semi-specific ISJ markers are based on sequences DNA was precipitated and dissolved in sterile, that are common in plants and partly comple- deionized H2O. The purity of DNA samples was mentary to the sequences at the intron–exon assessed spectrophotometrically and it reached splice junction (Rafalski 1997). The sequences 90%–95%. The DNA content of the samples was of ISJ primers used for DNA amplification in from 50.4 µg to 506.1 µg. this study are given in Table 2. Table 1. List of analyzed Polygonatum populations from Poland.
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