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Ribosomal, Telomeric and Heterochromatin Sequences Localization in the Karyotype of Anemone Hortensis

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Ribosomal, Telomeric and Heterochromatin Sequences Localization in the Karyotype of Anemone Hortensis ET AL . Botanical Journal of the Linnean Society, 2006, 150, 177–186. With 14 figures Ribosomal, telomeric and heterochromatin sequences localization in the karyotype of Anemone hortensis JELENA MLINAREC, DRAXEN A. PAPET and VITNJA BESENDORFER* Downloaded from https://academic.oup.com/botlinnean/article/150/2/177/2420480 by guest on 30 September 2021 Department of Molecular Biology, Division of Biology, Faculty of Science, University of Zagreb, Rooseveltov trg 6, HR-10000 Zagreb, Croatia Received February 2005; accepted for publication July 2005 The karyotype of the Mediterranean species Anemone hortensis L. (Ranunculaceae) was characterized with empha- sis on heterochromatin distribution and localization of ribosomal (18S−5.8S−26S and 5S rDNA) and telomeric repeats (TTTAGGG). Diploid chromosome complement, 2n = 2x = 16, common to all investigated populations, con- sisted of three acrocentric, one meta-submetacentric and four metacentric chromosomes ranging in size from 6.34 to 10.47 µm. Fluorescence in situ hybridization (FISH) with 18S and 5S rDNA probes revealed two 18S−5.8S−26S rDNA loci on a satellite and secondary constriction of acrocentric chromosome pair 2 and terminally on acrocentric chromosome pair 3, and two 5S rDNA loci in the pericentromeric region of meta-submetacentric chromosome pair 4 and in the proximity of the 18S−5.8S−26S rDNA locus on chromosome pair 2. The only GC-rich heterochromatin, as revealed by fluorochrome Chromomycin A3 staining, was that associated with nucleolar organizer regions, whereas AT-rich heterochromatin, stained with 4,6-diamino-2-phenylindole (DAPI), was distributed intercalarly and termi- nally on the long arm of all three acrocentric chromosomes, and terminally on chromosomes 4 and 5. FISH with Ara- bidopsis-type telomeric repeats (TTTAGGG) as a probe revealed two classes of signals, small dot-like and large bands, at chromosome termini exclusively, where they corresponded to terminal DAPI-stained heterochromatin. Het- eromorphism of chromosome pair 4, which refers to terminal DAPI bands and FISH signals, was observed in pop- ulations of Anemone hortensis. Chromosome pairing during meiosis was regular with formation of localized chiasmata proximal to the centromere. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 150, 177–186. ADDITIONAL KEYWORDS: fluorescence in situ hybridization – fluorochrome banding – karyotype analysis – meiosis – nucleolar organizer region – rRNA genes – telomeric repeats. INTRODUCTION 1995; Ehrendorfer & Samuel, 2001; Schuettpelz et al., 2002). According to these data, A. hortensis is placed The genus Anemone (Ranunculaceae) comprises 70–90 in the Coronaria group together with all tuberous species of perennial, low-growing herbs. Two subgen- anemones from the Mediterranean region and south- era are recognized in the genus, which correlate with central USA and one disjunct species from South base chromosome number x = 7 for the subgenus America. Anemonidium and x = 8 for the subgenus Anemone. Species of Anemone were considered favourable Anemone hortensis L. as a tuberous Mediterranean plant material for cytogenetic studies because of species belonging to the subgenus Anemone is wide- chromosomal polymorphism, high levels of hetero- spread all along the Croatian Adriatic coast. Consid- chromatin and variation in its distribution, meiotic erable efforts, based on morphological and molecular chromosome behaviour as well as variation in total data, have been made in recent decades to improve our DNA content between species (Böchner, 1945; Rothfels knowledge of the phylogenetic relationships of the spe- et al., 1966; Baumberger, 1970; Cullis & Schweizer, cies of Anemone (Hoot, Reznicek & Palmer, 1994; Hoot, 1974; Marks & Schweizer, 1974). Cytogenetic analysis of anemones has mainly focused on determination of chromosome morphology and heterochromatin distri- *Corresponding author. E-mail: [email protected] bution using the Giemsa C-banding technique (Marks © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 150, 177–186 177 178 J. MLINAREC ET AL. & Schweizer, 1974). Cytogenetically, A. blanda has with the fluorochromes Chromomycin A3 (CMA) and been the most extensively explored species owing to its DAPI and FISH mapping of the ribosomal and telom- unique karyotype among anemones; levels and distri- eric sequences were employed. In addition, the behav- bution of heterochromatin have been well described iour of nucleolar organizer regions (NORs) in mitosis (Marks & Schweizer, 1974; Marks, 1976). Due to a and meiosis was analysed. large proportion (53–67%) of repetitive DNA found in the genome, efforts have been made to characterize MATERIAL AND METHODS satellite DNAs, as constitutive parts of heterochroma- tin, and determine their chromosomal position in PLANT MATERIAL A. blanda by fluorescence in situ hybridization (FISH) Plants forming part of the natural populations of the Downloaded from https://academic.oup.com/botlinnean/article/150/2/177/2420480 by guest on 30 September 2021 (Cullis & Schweizer, 1974; Hagemann, Scheer & Adriatic islands of Hvar, Murter, Tisno, Krk and Vis, Schweizer, 1993). In A. blanda, a tandemly repetitive and mainland regions Makarska-Batko polje, Makar- sequence family (AbS1) is found to be located in all ska-Dugit, Makarska-Osejava and Starigrad-Paklen- 4,6-diamino-2-phenylindole (DAPI)-positive interca- ica were collected and potted in the Botanical Garden, lary bands and in the terminal DAPI-positive band Department of Botany, Faculty of Science, University of chromosome 3, whereas a dispersed repeated of Zagreb (Fig. 1). sequence (Hd), a part of the larger dorf-1 element that exhibits partial homology to the Lilium gypsy-type ele- ment del1, is preferentially associated with euchro- CHROMOSOME PREPARATION matic chromosome regions (Hagemann et al., 1993). Root tips were pretreated with 0.05% (w/v) colchicine Telomeres protect chromosome ends from degrada- (Sigma) at room temperature for 4 h, fixed in 3 : 1 (v/ tion and gradual shortening, maintain their structure v) ethanol/acetic acid at 4 °C for 24–48 h and stored in and regular segregation during the cell cycle, and 70% (v/v) ethanol. Floral buds collected in the field thereby contribute to genome stability. Telomeres were fixed directly in 3 : 1 (v/v) ethanol/acetic acid. consist of telomeric repeats, which are remarkably For karyotype observation, chromosomes were conserved in eukaryotes. The majority of plant stained with 1% (w/v) acetocarmine. Chromosome species analysed thus far possess Arabidopsis-type preparations used for fluorochrome and silver staining TTTAGGG telomeric repeats. However, a number of as well as in FISH experiments were prepared either exceptions have been described. Some species in the by softening in a mixture of 3% cellulase (Onozuka) order Asparagales (e.g. Allium) and in the family and 20% pectinase (Sigma) in 0.01 M citric buffer at Solanaceae (e.g. Cestrum) have been found to lack 37 °C for 45–55 min or by staining in 1% (w/v) aceto- Arabidopsis-type sequence repeats (Fuchs, Brandes & carmin, followed by maceration in 45% (v/v) acetic Schubert, 1995; Sykorová et al., 2003). Recently, inves- acid. A coverslip was removed from a slide by using tigations of chromosome ends in Aloe (Asphodelaceae), the dry-ice method (Sharma & Sharma, 1972) and air- Othocallis and Hyacinthella (Hyacinthaceae), all dried at room temperature for several days. belonging to the Asparagales, have revealed the pres- In the study of meiotic events, pollen mother cells ence of vertebrate-type TTAGGG telomeric repeats (PMCs) were extracted from anthers and slightly (Weiss & Scherthan, 2002; Puizina et al., 2003; Weiss- flamed in 45% acetic acid. Chromosomes in different Schneeweiss et al., 2004). As far as we are aware, stages of prophase I that were outside the cell walls there are no data regarding the characteristics of were selected for FISH experiments. telomeres in Ranunculaceae, including the genus Anemone. The most widely spread Mediterranean anemone in FLUOROCHROME AND SILVER STAINING Croatia is Anemone hortensis, which has not yet been Fluorochrome staining with CMA and DAPI was per- cytogenetically investigated. The only morphological formed according to the protocol of Kondo & Hizume study of A. hortensis in Croatia was by Radid (1987) on (1982) with slight modification. Slides were stained populations from Biokovo Mountain and the Makar- with 0.1 mg L−1 CMA for 15 min and counterstained ska region. Radid’s study showed that A. hortensis with 0.1% (v/v) methyl green or with 2 µg L−1 DAPI for appeared in a number of variants, such as A. hortensis 10 min. Silver staining was performed with colloidal var. heldreichii and A. hortensis var. stellata. The aim developer according to the method of Howell & Black here is to analyse the karyotype structure and genome (1980). organization of this species, in natural island and mainland populations including those described by Radid (1987), using morphometric data, the distribu- FLUORESCENCE IN SITU HYBRIDIZATION tion of heterochromatin, and the position of 5S and The position and number of 5S rDNA sites was 18S−5.8S−26S rDNA loci. For that purpose, staining determined by FISH, according to the method of © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 150, 177–186 GENOME ORGANIZATION OF ANEMONE HORTENSIS 179 I. Downloaded from https://academic.oup.com/botlinnean/article/150/2/177/2420480 by guest on 30 September 2021 II.
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