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The DNA Weights Per Nucleus (Genome Size) of More Than 2350 Species of the Flora of the Netherlands, of Which 1370 Are New to Sc

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The DNA Weights Per Nucleus (Genome Size) of More Than 2350 Species of the Flora of the Netherlands, of Which 1370 Are New to Sc 24 Forum geobotanicum (2019) 8: 24−78 DOI 10.3264/FG.2019.1022 Ben J.M. Zonneveld The DNA weights per nucleus (genome size) of more than 2350 species of the Flora of The Netherlands, of which 1370 are new to science, including the pattern of their DNA peaks Published online: 22 October 2019 © Forum geobotanicum 2019 Abstract Besides external characteristics and reading a piece were previously measured completely in most cases (‘t Hart et of DNA (barcode), the DNA weight per nucleus (genome size) al. 2003: Veldkamp and Zonneveld 2011; Soes et al. 2012; via flow cytometry is a key value to detect species and hybrids Dirkse et al. 2014, 2015; Verloove et al. 2017; Zonneveld [et and determine ploidy. In addition, the DNA weight appears to al.] 2000−2018), it can be noted that even if all species of a be related to various properties, such as the size of the cell and genus have the same number of chromosomes, there can still the nucleus, the duration of mitosis and meiosis and the be a difference of up to three times in the weight of the DNA. generation time. Sometimes it is even possible to distinguish Therefore, a twice larger DNA weight does not have to indicate between groups or sections, which can lead to new four sets of chromosomes. Finally, this research has also found classification of the genera. The variation in DNA weight is clues to examine further the current taxonomy of a number of also useful to analyze biodiversity, genome evolution and species or genera. relationships between related taxa. Moreover, it is important to know how large a genome is before one determines the base Samenvatting Naast uiterlijke kenmerken en het lezen van een sequence of the DNA of a plant. Flow cytometry is also stukje DNA (barcodering), is het DNA-gewicht per kern (via important for understanding fundamental processes in plants flow cytometrie) een sleutelwaarde om soorten en hybriden te such as growth and development and recognizing chimeras. In detecteren en ploïdie te bepalen. Daarnaast blijkt het DNA- the literature, DNA weight measurements are usually limited gewicht met verschillende eigenschappen samen te hangen, to one genus and often only locally (Siljak et al. 2010; Bai et zoals met de grootte van de cel en de kern, de duur van de al. 2012). In this study, however, it was decided to investigate mitose en meiose en met de generatie tijd. Soms is het zelfs all vascular plants from one country. This can also contribute mogelijk om tussen groepen of secties onderscheid te maken, to the protection of rare plants. This study is the first flora in wat tot nieuwe indeling van de geslachten kan leiden. De the world whose weight of DNA per nucleus and peak patterns variatie in DNA-gewicht is ook nuttig om biodiversiteit, has been determined. More than 6400 plants, representing genoom evolutie en relaties tussen verwante taxa te analyseren. more than 2350 (sub)species (more than 90%) have been Bovendien is van belang te weten, hoe groot een genoom is, collected, thanks to the help of almost 100 volunteers of alvorens men van een plant de base-volgorde van het hele Floristisch Onderzoek Nederland (Floron). Multiple DNA kan bepalen. Flow cytometrie is ook belangrijk voor het specimens of many species have therefore been measured, begrijpen van fundamentele processen in planten zoals groei preferably from different populations, in some cases more than en ontwikkeling en het herkennen van chimaeren. In de fifty. For 1370 species, these values were not previously literatuur wordt het meten van het DNA-gewicht meestal published. Moreover, a good number of the remaining 45% are beperkt tot één geslacht en ook nog vaak alleen lokaal. In deze new for The Netherlands. In principle, each species has a fixed studie is echter besloten om alle vaatplanten van één land te weight of DNA per nucleus. It has also been found that, onderzoeken. Dit kan dan ook een bijdrage leveren bij de especially between the genera, there are strong differences in bescherming van zeldzame planten. Deze studie is de eerste the number of peaks that determine the DNA weight, from one flora ter wereld waarvan het gewicht van het DNA per kern en to five peaks. This indicates that in a plant or organ there are de piekpatronen zijn bepaald. Hiervoor zijn ruim 6400 planten sometimes nuclei with multiples of its standard DNA weight verzameld die ruim 2350 (onder)soorten (meer dan 90 %) (multiple ploidy levels). It is impossible to show graphs of vertegenwoordigen, vooral dankzij de hulp van bijna 100 more than 2350 species. Therefore, we have chosen to show vrijwilligers van Floristisch Onderzoek Nederland (Floron). the peak pattern in a new way in a short formula. Within most Van veel soorten zijn dus meerdere exemplaren gemeten, bij genera there are clear differences in the DNA weights per voorkeur uit verschillende populaties, van sommigen wel meer nucleus between the species, in some other genera the DNA dan vijftig. Voor 1370 soorten zijn dit niet eerder weight is hardly variable. Based on about twenty genera that gepubliceerde waarden. Bovendien is van de overige 45 % ook 25 een flink aantal nieuw voor Nederland. In principe heeft elke Plants and animals are made up of cells. Each cell contains a soort een vast DNA- gewicht per kern. Er is ook gevonden dat nucleus. Each nucleus contains long double strands of DNA vooral tussen geslachten, er sterke verschillen zijn in het aantal made up of four bases, abbreviated with A, T, G and C. These pieken die het DNA-gewicht aangeven, van een tot wel vijf strands are surrounded by proteins. DNA strands + proteins are pieken. Dit geeft aan dat in een plant of orgaan er soms kernen divided into a number of pieces that are called chromosomes. zijn met veelvouden van het standaard DNA-gewicht Chromosomes can vary greatly in size and number in different (meerdere ploïdie niveaus). Het is ondoenlijk om grafieken van organisms, so the number of chromosomes is not directly a alle 2350 soorten weer te geven. Daarom is er voor gekozen measure of the weight of DNA in the nucleus. In diploid om het piekenpatroon op een nieuwe manier in een korte organisms, each chromosome is present in duplicate, so there formule weer te geven. Binnen de meeste geslachten zijn er are two sets of chromosomes. In most organisms only a limited duidelijk verschillen in de DNA-gewichten per kern tussen de part of this DNA codes for genes or characteristics. In humans soorten, in enkele andere geslachten is het DNA-gewicht with 7 picograms (pg) of DNA per nucleus, this is less than nauwelijks variabel. Gebaseerd op een twintigtal geslachten 2%, representing about 25.000 genes. die eerder in de meeste gevallen compleet gemeten zijn, kan A number of alternative, not entirely overlapping terms are opgemerkt worden dat ook als alle soorten van een geslacht used for the weight of DNA per nucleus: weight of DNA per hetzelfde aantal chromosomen hebben, er toch een verschil tot nucleus, quantity of DNA per nucleus, nuclear DNA weight, meer dan driemaal in het gewicht van het DNA kan zijn. Een 2C-value, nuclear weight, genome size and genome weight. groter DNA-gewicht hoeft dus niet op meer dan 2 sets See 2x, 2n and 2C below for an explanation. It should be noted chromosomen te wijzen. Tenslotte heeft dit onde rzoek ook that nuclear weight is incorrect because the DNA represents aanwijzingen gevonden om de huidige taxonomie van een less than 1 permille of the nuclear weight. Here we use the aantal soorten of genera nader tegen het licht te houden. DNA weight or weight of the DNA or just DNA, whereby per nucleus and nuclear are often omitted. A code (e.g. 2C 4C 8C) Key words: Genome size, DNA weight, 2C-value, Dutch has been devised to indicate the number of peaks as seen in the Flora, peak profiles flow cytometer, with the highest peak in bold. These peaks represent the different number of chromosome sets (ploidies) measured in a sample. Genomic weight is actually 1C, so one set of chromosomes, but is pars pro toto also often used for Dr. Ben J. M. Zonneveld 2C-value. Kew DNA C-values (Bennet and Leitch 2012) is a NBC Naturalis collection of all published DNA weights of mainly Darwinweg 2 economically important crops with 8700 measurements. It 2333 CR Leiden, The Netherlands includes many duplicates and measurements with methods [email protected] other than propidium iodide. The DNA weight can be used for all kinds of applications, especially because it can be determined quickly and relatively cheaply. Commercially, it is usually used to check whether a plant is (has become) Abbreviations Aa/aa: main standard, only mentioned in polyploid or to determine the number of bases of the nuclear special cases; Aa15.9: Agave americana 'Aureomarginata' DNA before sequencing. 15.9 pg; ad: adventive; A'dam: Amsterdam; Afrik. Pl.: In this article we want to give an overview of the DNA weights Afrikaander plein; AGS: Alpine garden society UK; arb.: of the Dutch flora, but also reflect on the following questions: arboretum; BG: Botanical Garden; BZ: B. Zonneveld added; Are we dealing with one or two species? Are there differences c.: circa, about; CCDB: Chromosome Counts Database; in DNA weight within or between genera? Does the weight of Chl22.8: Chlorophytum comosum 22.8 pg; cu: cultivated; E: the DNA mean that the right species is measured? What is the East; ex: exote; FL: Floron list recognized after 2005; gr: possible other parent of this hybrid? How do the DNA weights groot/large; haem72: Haemanthus albiflos 72pg; hort.: relate between plants with different growth habits and hortorum/from the garden; HT: Heemtuin; k.: kade; kl: environments? What is the relationship between chromosome klein/small; l.: lane; L.dorp: Leiderdorp; N: North; NDV: Ned.
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