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We present a phylogenetic dating of asterids, based on a taxon tree representing all major groups and orders and 83 of the families of asterids, with an underlying data set comprising six chloroplast DNA markers totaling positions. Phylogenetic dating was done with bhm dating Lafayette rate smoothing by penalized likelihood. Confidence intervals were calculated by bootstrapping. Six reference fossils were used for calibration.

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The genus Rosa comprises more than woody species characterized by intensive hybridization, introgression, and an overall complex evolutionary history. Here we analyzed 5S ribosomal DNA in 19 species covering two subgenera and the major sections within subg.

In addition to diploids and polyploids with regular meiosis, we focused on 5x dogroses Rosa sect. Caninaewhich exhibit an asymmetric meiosis differentiating between bivalent- and univalent-forming chromosomes. Using genomic resources, we reconstructed 5S rDNA units to reveal their phylogenetic relationships.

New dating in Killeen TX, we deed locus-specific probes derived from intergenic spacers IGSs and determined the position and of 5S rDNA families on chromosomes. Within subg.

Associated data

Rosa species of sect. The allo-pentaploid dogroses showed a distinct distribution of 5S rDNA families between bivalent- and univalent-forming chromosomes.

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In conclusion, two divergent 5S rDNA families dominate rose genomes. Both gene families apparently arose in the early history of the genus, already 30 myrs ago, and apparently survived numerous speciation events thereafter.

These observations are consistent with a relatively slow genome turnover in the Rosa genus. Each unit consists of an evolutionary conserved coding region of bp and a variable intergenic spacer Dating with Missouri ladies Long and Dawid, The units within the 5S arrays retain a high degree of identity due to homogenizing forces referred to as concerted evolution Dover, ; Eickbush and Eickbush, where unequal crossing-over and gene conversion are major forces driving the process.

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Regardless of the mechanism, numerous factors such as the of arrays, their mutation rate, formation of new variants, the intensity of natural selection, or dating Vermont aged men population size can affect homogenization of repeats Dover, ; Ohta, ; Nagylaki, As a consequence, two or more variants differing in the length and nucleotide sequence may simultaneously exist per genome Cronn et al. The genus Rosa L. Rosaceae comprises about species widely distributed across the northern hemisphere.

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Taxonomy is considered to be challenging because frequent polyploidy in app. The existence of multiple cytotypes and variable degree of retention of progenitor alleles leading to incomplete lineage sorting complicating taxonomic classifications.

Data and methods

In addition, species identification is generally hampered because most species are characterized rather by combinations of morphological characters than by single discriminating traits Christ, ; Wissemann, Moreover, roses are one of oldest ornamentals Wang, and their complex history of cultivation and breeding may generate another uncertainty in phylogenetic studies.

Several attempts have been made to reconstruct the phylogeny of the genus Millan et al. Currently, the system comprises four subgenera: Hulthemia one speciesHesperhodos two speciesPlatyrhodon one speciesand Rosamike Myrtle dating latter consisting of 10 sections free chatroom Mobile AL comprising the vast majority of species Wissemann, The most recent phylogenies detected Rosa persica subg. Hulthemia and Rosa minutifolia subg.

Hesperhodos as early diverging lineages, and a major split of the speed dating Baltimore Maryland MD gratis into two large clades: the Synstylae and allies clade consisting of sect. The exclusively polyploid members of a large section Caninae DC. Canina meiosis in a strongly matroclinal inheritance of genetic information since two pairing genomes form bivalents, while the remaining genomes remain unpaired as univalents and are transmitted by the female germ line only.

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Amazingly, in plastid phylogenies, sect. Caninae appeared to be polyphyletic since species with fragrant glands subsect.

Original research article

Rubigineae and Vestitae were separated from the remaining species subsect. Caninae by Rosa gallica and Rosa arvensis which perform regular meiosis Wissemann and Ritz, ; Fougere-Danezan et al. Thus, Canina meiosis has been probably evolved twice, which is supported by fluorescence in situ hybridization FISH analyses of meiotic chromosomes Herklotz et al. Ribosomal DNA loci have been studied in several diploid free trial phone chat Atlanta Georgia GA polyploid species of Rosa so far.

Ma et al. Fernandez-Romero et al. In pentaploid dogroses, four to five 35S loci were reported implying the occasional loss of one locus Lim et al. The 5S locus has been less commonly studied, while there is evidence for more than one 5S locus per haploid set.

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The mature women dating Macon of 5S rDNA clones from diploid Rosa rugosa revealed a conserved bipartite polymerase III promoter and non-coding IGS region Tynkevich and Volkov, b evidencing that organization at the unit level is similar to most other plants. In contrast, the level of IGS similarity between R. In this study, based on genomic and cytogenetic approaches, we aim to map the evolutionary history of 5S rDNA loci across the genus Rosa. Bioinformatic methods were used to determine the abundance and homogeneity of 5S rDNA in the genomes.

Table 1. List of Rosaceae species used in this study, ploidy, source, and read archive accessions and the analyses employed.

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Material of polyploid dogroses was sampled in wild populations in Germany and the Czechia Supplementary Table S1. Diploid species and tetraploid species with regular meiosis were obtained from various Botanical Gardens Supplementary Table S1. In addition, we retrieved sequence information from published work stored in the ENA database for bioinformatics analyses Supplementary Table S1.

Total genomic DNA dating in the dark Lakewood CO online Rosa canina was extracted from fresh leaves applying the standard protocol Rogers and Bendich, The length of amplified products was nt for the A variant and nt for the B variant.

For bioinformatic analyses, the whole genomic sequencing data for 19 Rosa accessions were used Supplementary Table S2. For the more distantly related species, Rosa spinosissima and R. The genome abundance and copy was calculated from genome proportions according to the formula stated in Lunerova et al.

Figure 1. B Dot plot of pairwise comparisons clones. Note that only coding regions showed ificant similarity. This resulted in two re out of The two re were aligned and served in a second round with same parameters as mapping scaffold of bp length. In the second mapping, 50 re out of Based on this model, we computed a maximum-likelihood tree in MEGA X whose branch support was evaluated by bootstrap replicates.

Rooted with G. Therefore, we used two calibration points along the tree taken from the Atlas PA date ideas fossils given in Xiang et al. A timetree inferred by applying the RelTime method Tamura et al.

Materials and methods

All positions containing gaps and missing data were eliminated complete deletion option. A neighbor ing tree was constructed with the Naperville IL women free dating software Gouy et al.

The fastq re were initially filtered for quality and trimmed to uniform length using the pre-processing and QC tools in RepeatExplorer2 Novak et al. Read length ranged between and bp, depending on sequencing library and the Illumina sequencing platform. We used 1 million paired-end re, or 1 million single-end re as inputs for RepeatExplorer2 clustering.

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This bioinformatic pipeline runs a graph-based clustering algorithm Novak et al. The similarity and structure-based repeat identification tools in RepeatExplorer2 aid in identification of the repeats. We also used the SeqGrapheR Novak et al. For slide preparations, we used young anthers from flower buds of dating agency Fayetteville 0. Before slide preparation, anthers were pre-treated by 0.

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FISH teen dating College Station 18 the procedures described in Herklotz et al. Slide preparation and hybridization followed standard protocols Schwarzacher and Heslop-Harrison, The slides were scanned using epifluorescence microscopes Olympus Provis AX70, with cold cube camera, Metasystems, Germany.

Sequence analysis revealed some conserved regulatory elements: Box-C within the coding region, the TATA box at —29 both clonesand T-rich terminators downstream of the coding sequences Figure 1A.

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Box-A could not be unambiguously determined due to primer overlap. Intragenomic homogeneity was high, and no ificant SNPs were revealed in mapping experiments not shown. Pairwise alignment revealed conserved coding regions, while most of the IGS was dissimilar between both sequences Figure 1B.

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We took advantage of considerable sequence divergence between both clones and amplified the locus-specific IGS subregions from plasmids. To determine the abundance of individual 5S rRNA gene families in Rosa genomes, we used available genomic resources Table 1.

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Both families appeared to be equally homogeneous containing a relatively low of SNPs consistent with our findings obtained by comparison of individual 5S rDNA clones Tynkevich and Volkov, ab.