{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T15:41:13Z","timestamp":1784130073327,"version":"3.55.0"},"reference-count":61,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2021,5,13]],"date-time":"2021-05-13T00:00:00Z","timestamp":1620864000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Breeding programs in ornamentals can be facilitated by integrating knowledge of phylogenetic relatedness of potential parents along with other genomic information. Using AFLP, genetic distances were determined for 59 Geranium genotypes, comprising 55 commercial cultivars of the three subgenera of a total collection of 61 Geranium genotypes. A subgroup of 45 genotypes, including intragroup and intergroup hybrids, were selected and further characterized for genome sizes and chromosome numbers. The variation in genome size ranged from 1.51 \u00b1 0.01 pg\/2C to 12.94 \u00b1 0.07 pg\/2C. The chromosome numbers ranged from 26 to 108\u2013110 with some hybrids showing an aberrant number of chromosomes based on their parents\u2019 constitution. All chromosome numbers of Geranium are an even number, which presumes that unreduced gametes occur in some cross combinations. Overall, parental difference in genome size and chromosome number were not limiting for cross compatibility. Good crossing compatibility was correlated to a Jaccard similarity coefficient as parameter for parental relatedness of about 0.5. Additionally, parent combinations with high differences in the DNA\/chromosome value could not result in a successful cross. We expect that our results will enable breeding programs to overcome crossing barriers and support further breeding initiatives.<\/jats:p>","DOI":"10.3390\/genes12050730","type":"journal-article","created":{"date-parts":[[2021,5,13]],"date-time":"2021-05-13T11:10:06Z","timestamp":1620904206000},"page":"730","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":18,"title":["Can Knowledge of Genetic Distances, Genome Sizes and Chromosome Numbers Support Breeding Programs in Hardy Geraniums?"],"prefix":"10.3390","volume":"12","author":[{"given":"Mehrdad","family":"Akbarzadeh","sequence":"first","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"},{"name":"Department Plants and Crops, Faculty of Bioscience Engineering, Ghent University, Coupure Links 653, 9000 Ghent, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5398-5489","authenticated-orcid":false,"given":"Katrijn","family":"Van Laere","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8703-3651","authenticated-orcid":false,"given":"Leen","family":"Leus","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"De Riek","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5398-8257","authenticated-orcid":false,"given":"Johan","family":"Van Huylenbroeck","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefaan P.O.","family":"Werbrouck","sequence":"additional","affiliation":[{"name":"Department Plants and Crops, Faculty of Bioscience Engineering, Ghent University, Coupure Links 653, 9000 Ghent, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8497-1220","authenticated-orcid":false,"given":"Emmy","family":"Dhooghe","sequence":"additional","affiliation":[{"name":"Plant Sciences Unit, Flanders Research Institute for Agricultural, Fisheries and Food Research (ILVO), Caritasstraat 39, 9090 Melle, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,5,13]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"594","DOI":"10.2307\/2992018","article-title":"Taxonomic revision of Geranium sections batrachioidea and divaricata (Geraniaceae)","volume":"85","author":"Aedo","year":"1998","journal-title":"Ann. Missouri Bot. Gard."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1600\/0363644054782260","article-title":"Taxonomic revision of Geranium sect. Dissecta (Geraniaceae)","volume":"30","author":"Aedo","year":"2005","journal-title":"Syst. Bot."},{"key":"ref_3","first-page":"157","article-title":"Geraniaceae","volume":"Volume 5","author":"Kubitzki","year":"2007","journal-title":"Flowering Plants\u00b7Eudicots"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"661","DOI":"10.2307\/2399852","article-title":"Phylogenetic Relationships of the Geraniaceae and Geraniales from rbcL Sequence Comparisons","volume":"80","author":"Price","year":"1993","journal-title":"Ann. Missouri Bot. Gard."},{"key":"ref_5","first-page":"205","article-title":"Taxonomic revision of Geranium sect. Brasiliensia (Geraniaceae)","volume":"26","author":"Aedo","year":"2001","journal-title":"Syst. Bot."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1080\/00288250909509789","article-title":"Phylogenetic relationships of Geranium species indigenous to New Zealand","volume":"47","author":"Mitchell","year":"2009","journal-title":"N. Z. J. Bot."},{"key":"ref_7","unstructured":"Yeo, P.F. (2001). Hardy Geraniums, Timber Press, Inc.. [2rd ed.]."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1016\/j.ympev.2017.03.012","article-title":"Species-level phylogeny, fruit evolution and diversification history of Geranium (Geraniaceae)","volume":"110","author":"Marcussen","year":"2017","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_9","first-page":"1","article-title":"RHS Plant Trials and Awards Hardy Geraniums\u2014Stage 3 and Geraniums Suitable for Rock Gardens Stage 3 of the Trial of Hardy Geraniums and Geraniums Suitable for Rock Gardens","volume":"17","author":"Armitage","year":"2007","journal-title":"RHS"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1093\/aob\/mct185","article-title":"Nutrient reserves may allow for genome size increase: Evidence from comparison of geophytes and their sister non-geophytic relatives","volume":"112","year":"2013","journal-title":"Ann. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1093\/aob\/mcr267","article-title":"Genome size and DNA base composition of geophytes: The mirror of phenology and ecology?","volume":"109","year":"2012","journal-title":"Ann. Bot."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1093\/aob\/mch151","article-title":"Genome size variation in Central European species of Cirsium (Compositae) and their natural hybrids","volume":"94","author":"Wang","year":"2004","journal-title":"Ann. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1139\/G09-056","article-title":"Genome size variation in Begonia","volume":"52","author":"Dewitte","year":"2009","journal-title":"Genome"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1166\/asl.2010.1115","article-title":"Towards a genome size and chromosome number database of balkan flora: C-values in 343 taxa with novel values for 242","volume":"3","author":"Pustahija","year":"2010","journal-title":"Adv. Sci. Lett."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1756-1051.2010.01028.x","article-title":"Geranium kikianum sp. nov. (Geraniaceae) from the southern Peloponnese, Greece","volume":"29","author":"Tan","year":"2011","journal-title":"Nord. J. Bot."},{"key":"ref_16","first-page":"24","article-title":"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","volume":"8","author":"Zonneveld","year":"2019","journal-title":"Forum Geobot."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1111\/j.1095-8339.2003.00258.x","article-title":"The morphology and affinities of Geranium sections Lucida and Unguiculata","volume":"144","author":"Yeo","year":"2004","journal-title":"Bot. J. Linn. Soc."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s13237-019-00299-8","article-title":"A case of intraspecific euploidy (2\u00d7, 4\u00d7) and secondary chromosomal associations in wild accessions of Geranium pratense L. from cold deserts of Ladakh (India)","volume":"63","author":"Khan","year":"2020","journal-title":"Nucleus"},{"key":"ref_19","first-page":"39","article-title":"The genus geranium L. (geraniaceae) in North America. I. Annual species","volume":"58","author":"Aedo","year":"2000","journal-title":"An. Jard\u00edn Bot\u00e1nico Madr."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"72","DOI":"10.2307\/2445884","article-title":"Phylogenetic position of the Hawaiian geraniums based on rbc L Sequences","volume":"84","author":"Pax","year":"1997","journal-title":"Am. J. Bot."},{"key":"ref_21","first-page":"5467","article-title":"Phylogenetic relationships of geranium species in different regions of Turkey","volume":"25","author":"Koloren","year":"2016","journal-title":"Fresenius Environ. Bull."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"326","DOI":"10.1600\/036364408784571482","article-title":"Phylogeny and historical biogeography of geraniaceae in relation to climate changes and pollination ecology","volume":"33","author":"Fiz","year":"2008","journal-title":"Syst. Bot."},{"key":"ref_23","first-page":"161","article-title":"Bridging the gap: Tools for interspecific and intergeneric hybridization in ornamentals","volume":"1283","author":"Eeckhaut","year":"2020","journal-title":"Acta Hortic."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Kubota, S., Konno, I., and Kanno, A. (2012). Molecular phylogeny of the genus Asparagus (Asparagaceae) explains interspecific crossability between the garden asparagus (A. officinalis) and other Asparagus species. Theor. Appl. Genet.","DOI":"10.1007\/s00122-011-1709-2"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"625","DOI":"10.2503\/jjshs.65.625","article-title":"Cross compatibility of Petunia cultivars and P. axillaris with native taxa of Petunia in relation to their chromosome number","volume":"65","author":"Watanabe","year":"1996","journal-title":"J. Jpn. Soc. Hortic. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1007\/s10681-008-9844-9","article-title":"Interspecific hybridisation and genome size analysis in Buddleja","volume":"166","author":"Leus","year":"2009","journal-title":"Euphytica"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1007\/s10681-017-1934-0","article-title":"Interspecific hybridization in Sarcococca supported by analysis of ploidy level, genome size and genetic relationships","volume":"213","author":"Denaeghel","year":"2017","journal-title":"Euphytica"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Van Huylenbroeck, J. (2018). Helleborus. Ornamental Crops, Springer International Publishing.","DOI":"10.1007\/978-3-319-90698-0"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1534","DOI":"10.21273\/HORTSCI.44.6.1534","article-title":"Production and Verification of Hydrangea macrophylla \u00d7 H. angustipetala Hybrids","volume":"44","author":"Kardos","year":"2009","journal-title":"HortScience"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"803","DOI":"10.2503\/jjshs.68.803","article-title":"Production of interspecific hybrid plants through cotyledonary segment culture of embryos derived from crosses between Hydrangea macrophylla f. hortensia (Lam.) Rehd. and H. arborescens L","volume":"68","author":"Kudo","year":"1999","journal-title":"J. Jpn. Soc. Hortic. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"85","DOI":"10.24266\/0738-2898-19.2.85","article-title":"Verification and Establishment of Hydrangea macrophylla \u2018Kardinal\u2019 \u00d7 H. paniculata \u2018Brussels Lace\u2019 Interspecific Hybrids","volume":"19","author":"Reed","year":"2001","journal-title":"J. Environ. Hortic."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1600\/036364410792495827","article-title":"Unraveling extensive paraphyly in the genus Hydrangea s. l. with implications for the systematics of tribe Hydrangeeae","volume":"35","author":"Samain","year":"2010","journal-title":"Syst. Bot."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"84","DOI":"10.21273\/JASHS.133.1.84","article-title":"Production and characterization of intergeneric hybrids between Dichroa febrifuga and Hydrangea macrophylla","volume":"133","author":"Reed","year":"2008","journal-title":"J. Am. Soc. Hortic. Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s00606-011-0422-6","article-title":"Genetic relationships in European and Asiatic Buxus species based on AFLP markers, genome sizes and chromosome numbers","volume":"293","author":"Hermans","year":"2011","journal-title":"Plant Syst. Evol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1007\/978-3-642-83962-7_18","article-title":"DNA protocols for plants","volume":"Volume 57","author":"Doyle","year":"1991","journal-title":"Molecular Techniques in Taxonomy"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"4407","DOI":"10.1093\/nar\/23.21.4407","article-title":"AFLP: A new technique for DNA fingerprinting","volume":"23","author":"Vos","year":"1995","journal-title":"Nucleic Acids Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1155","DOI":"10.1007\/s001220051320","article-title":"Validation of criteria for the selection of AFLP markers to assess the genetic variation of a breeders\u2019 collection of evergreen azaleas","volume":"99","author":"Dendauw","year":"1999","journal-title":"Theor. Appl. Genet."},{"key":"ref_38","unstructured":"Elmer, P. (1995). AFLP Plant Mapping Kit Protocol (P\/N 402083), Perkin Elmer."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1126\/science.220.4601.1049","article-title":"Rapid flow cytometric analysis of the cell cycle in intact plant tissues","volume":"220","author":"Galbraith","year":"1983","journal-title":"Science"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1093\/aob\/mci005","article-title":"Plant DNA flow cytometry and estimation of nuclear genome size","volume":"95","year":"2005","journal-title":"Ann. Bot."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Dole\u017eel, J., Sgorbati, S., and Lucretti, S. (1992). Comparison of three DNA fluorochromes for flow cytometric estimation of nuclear DNA content in plants. Physiol. Plant.","DOI":"10.1111\/j.1399-3054.1992.tb04764.x"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Lysak, M.A., and Dolezel, J. (1998). Estimation of nuclear DNA content in Sesleria (Poaceae). Caryologia.","DOI":"10.1080\/00087114.1998.10589127"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Dole\u017eel, J., Dole\u017eelov\u00e1, M., and Nov\u00e1k, F.J. (1994). Flow cytometric estimation of nuclear DNA amount in diploid bananas (Musa acuminata and M. balbisiana). Biol. Plant.","DOI":"10.1007\/BF02920930"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1093\/aob\/mci019","article-title":"The origin, evolution and proposed stabilization of the terms \u201cgenome size\u201d and \u201cC-value\u201d to describe nuclear DNA contents","volume":"95","author":"Greilhuber","year":"2005","journal-title":"Ann. Bot."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Kirov, I., Divashuk, M., Van Laere, K., Soloviev, A., and Khrustaleva, L. (2014). An easy \u201csteamDrop\u201d method for high quality plant chromosome preparation. Mol. Cytogenet.","DOI":"10.1186\/1755-8166-7-21"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Kirov, I., Khrustaleva, L., Van Laere, K., Soloviev, A., Meeus, S., Romanov, D., and Fesenko, I. (2017). DRAWID: User-friendly java software for chromosome measurements and idiogram drawing. Comp. Cytogenet.","DOI":"10.3897\/compcytogen.v11i4.20830"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1095-8339.1984.tb00998.x","article-title":"Fruit-discharge-type in Geranium (Geraniaceae): Its use in classification and its evolutionary implications","volume":"89","author":"Yeo","year":"1984","journal-title":"Bot. J. Linn. Soc."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1007\/s00606-013-0924-5","article-title":"Cytological studies of Angiosperms (174 species) from District Kangra, Himachal Pradesh (India)","volume":"300","author":"Rani","year":"2013","journal-title":"Plant Syst. Evol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1508\/cytologia.81.183","article-title":"Chromosome counts through male meiosis in some dicots from the Hills around Shimla, Himachal Pradesh","volume":"81","author":"Singhal","year":"2016","journal-title":"Cytologia (Tokyo)"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1155\/2013\/248943","article-title":"Ethnobotany and Ethnomedicinal Uses, Chromosomal Status and Natural Propagation of Some Plants of Lahaul-Spiti and Adjoining Hills","volume":"2013","author":"Kumar","year":"2013","journal-title":"J. Bot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1007\/s00606-011-0516-1","article-title":"Chromosome number, male meiosis and pollen fertility in selected angiosperms of the cold deserts of Lahaul-Spiti and adjoining areas ( Himachal Pradesh, India )","volume":"297","author":"Kumar","year":"2011","journal-title":"Plant Syst. Evol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s13237-017-0211-x","article-title":"Intraspecific floral morphotypes in six high altitude perennial herbaceous species from northwest Himalaya: Their chromosome counts, meiotic behavior and pollen fertility","volume":"61","author":"Kumar","year":"2018","journal-title":"Nucleus"},{"key":"ref_53","first-page":"19","article-title":"Chromosome numbers of bulgarian angiosperms: An introduction to a chromosome atlas of the Bulgarian flora","volume":"3","author":"Kuzmanov","year":"1993","journal-title":"Flora Mediterr."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1080\/0028825X.2004.9512936","article-title":"Contributions to a chromosome atlas of the New Zealand flora\u201438. Counts for 50 families","volume":"42","author":"Murray","year":"2004","journal-title":"N. Z. J. Bot."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"353","DOI":"10.3372\/wi.39.39213","article-title":"Karyological data of some angiosperms from Romania","volume":"39","author":"Baltisberger","year":"2009","journal-title":"Willdenowia"},{"key":"ref_56","first-page":"93","article-title":"A cytological survey of eastern Himalayan plants. II","volume":"11","author":"Roy","year":"1988","journal-title":"Cell Chromosom. Res."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1007\/s11104-013-1794-x","article-title":"Small genomes dominate in plants growing on serpentine soils in West Balkans, an exhaustive study of 8 habitats covering 308 taxa","volume":"373","author":"Pustahija","year":"2013","journal-title":"Plant Soil"},{"key":"ref_58","first-page":"1","article-title":"Chemical composition and antioxidant activity of Geranium macrorrhizum in relation to ploidy level and environmental conditions","volume":"306","author":"Siljak","year":"2020","journal-title":"Plant Syst. Evol."},{"key":"ref_59","unstructured":"Freeman, L., Diener, J., Points, F., Ranch, R.R., and Points, F. (2011). United States Plant Patent. (US PP23,370 P2), U.S. Patent."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1080\/0028825X.2000.9512673","article-title":"Index of chromosome numbers of indigenous New Zealand spermatophytes","volume":"38","author":"Dawson","year":"2000","journal-title":"N. Z. J. Bot."},{"key":"ref_61","first-page":"13","article-title":"Interspecific hybridisation and polyploidisation as tools in ornamental plant breeding","volume":"612","author":"Lim","year":"2003","journal-title":"Acta Hortic."}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/5\/730\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:00:16Z","timestamp":1760162416000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/5\/730"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,5,13]]},"references-count":61,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2021,5]]}},"alternative-id":["genes12050730"],"URL":"https:\/\/doi.org\/10.3390\/genes12050730","relation":{},"ISSN":["2073-4425"],"issn-type":[{"value":"2073-4425","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,5,13]]}}}