Karyotype Traits in Grindelia Squarrosa (Pursh) Dunal (Asteraceae), an Invasive Plant in Romania
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Truta et. al.·Silvae Genetica (2012) 61-4/5, 179-186 Karyotype traits in Grindelia squarrosa (Pursh) Dunal (Asteraceae), an invasive plant in Romania By ELENA TRUTA1),*), GABRIELA VOCHITA1), ADRIAN OPREA2) and CULITA SIRBU3) (Received 3rd February 2012) Abstract Europe (Russia, the Ukraine, Republic of Moldova, Esto- The description of the karyotype features and idio- nia, Lithuania, Czech Republic, Belgium, Sweden, gram in Grindelia squarrosa (Pursh) Dunal (Aster- Latvia, Ireland) (SIRBU and OPREA, 2008). Although the aceae), an invasive plant in Romania, are reported here species is most common in the lower elevations of plains for the first time. The diploid chromosome number is and foothills, it was met, too, at 3000 m altitude in Col- 2n = 2x = 12, in agreement with the data published for orado and New Mexico. In Northern Utah, it occurs in the other species of the genus. The karyomorphological Tony Grove Canyon (2000 m) in the Cache National For- data show that the complements of the studied geno- est, also on disturbed ground throughout the valleys and types have small chromosomes (mean chromosome – in the adjacent Wasatch Mountains at elevations of at length is ± SE = 2.56 ± 0.10 µm, and mean length of X – least 2200 m (MCDONOUGH, 1975; WALSH, 1993). Report- haploid complements is ± SE = 15.33 ± 0.69 µm, with a X ed for the first time in the flora of Romania in 1998 range of variability comprised between 12.87–17.51 µm). The karyotypes are made up of six pairs of metacentric (SIRBU and OPREA, 1998), G. squarrosa can be considered and submetacentric chromosomes, with an identical an invasive alien plant in this country. Probably, the formula of the diploid complement: KF = 2n = 12 = 8m+ plant came into Romania by accident from the former 2sm + 2sm-SAT. Satellites are located on the short USSR, carried across by goods or passenger trains. It arms of the chromosomes of pair III. The karyotypes was first identified in the ruderal areas on the side- show a relatively high level of intra-specific uniformity tracks of the Socola-Iasi railway station. Meanwhile it as well as similar symmetry patterns (R = 1.29–1.53; invaded anthropic habitats both in the North-East and TF% = 38.78–41.57%; AsI %=54.54–57.61%; A1 = 0.24– the South of the Moldavian Region of Romania (SIRBU 0.32; A2 = 0.08–0.16), belonging to 1A and 2A classes of and OPREA, 2011). At present, G. squarrosa is considered symmetry. The small size of the chromosomes, the pres- as being fully naturalized in Romania, it having an evi- ence of only two chromosome morphometric types, and dent invasive tendency. Although in Romania the inva- the preponderance of metacentrics confer a relatively high degree of symmetry to the karyotypes studied. sion of this species into agricultural crops or native plant communities has not occurred yet, this trend is Key words: Grindelia squarrosa, invasive plant, karyotype, not excluded in the future, given its behaviour in the mitotic chromosomes, satellites, asymmetry indexes. neighbouring countries (SIRBU and OPREA, 2008, 2011). In the Ukraine and the Republic of Moldova, G. squar- Introduction rosa is considered as a very aggressive plant, while for Spain it is noted as potentially invasive (SANZ ELORZA Willd. is a genus with ca. 45 species in et Grindelia 2001). North America and 26 species in South America, com- al., prising annual, biennial, perennial forbs or subshrubs The invasive alien species, including curlycup widely distributed in xerophytic and halophytic areas gumweed, are seen as a major threat to the native biodi- (BAEZA and SCHRADER, 2005; DEBLE and OLIVEIRA-DEBLE, versity, ecosystem structure and conservation of the pro- 2010). In 1804, Grindelia seeds from Mexico were tected areas, thus causing damages to agriculture, brought to Europe (Royal Gardens – Madrid, Spain) and forestry, fisheries and other human activities, and cultivated as Aster spathulatus Hort.; afterwards they threatening human health (STINSON et al., 2006). Except were distributed to other botanical gardens (STEYER- the aspect of its invasiveness – sometimes seen as harm- MARK, 1937). ful – G. squarrosa is one of the only two officinal plants in the genus (the other is ) The curlycup gumweed – Grindelia squarrosa (Pursh) Grindelia G. robusta (GHEDIRA 2010). The chemical profile, represented Dunal (BRUMMITT and POWELL, 1992), a common weed et al., originating in the central prairies of North America is by diterpenes (grindelic acid and its methylesthers), now largely spread over Eastern, Central and Western flavonoids (quercetin, kempferol), tannins (5.3%), vita- min P, resin, phenolic acids, and essential oils (0.3–0.5%), confers numerous medical and pharmaceuti- 1) NIRDBS – Institute of Biological Research, Department of Cell cal valences to plant extracts and bio-preparations. Biology and Microbiology, 47 Lascar Catargi Street, 700107 Because of this complex chemical constitution, the Iasi, Romania. extracts are valuable as stimulants, sedatives, astrin- 2) Botanical Garden, 7–9 Dumbrava Rosie Street, 700487 Iasi, gents, purgatives, emetics, diuretics, antiseptics, and Romania. disinfectants and are used in the treatment of bronchial 3 ) University of Agricultural Sciences and Veterinary Medicine spasm, whooping cough, asthma, and rashes caused by Iasi, Faculty of Agriculture, 3 Mihail Sadoveanu Street, 700490 Iasi, Romania. poison ivy (Toxicodendron radicans L. Kunze), while the *) Corresponding author: ELENA TRUTA. Fax: +40-232218121. tinctures are useful for bladder and urethra infections E-mail: [email protected] (JOHNSON and NICHOLS, 1970; BARE, 1979). Silvae Genetica 61, 4–5 (2012) 179 DOI:10.1515/sg-2012-0023 edited by Thünen Institute of Forest Genetics Truta et. al.·Silvae Genetica (2012) 61-4/5, 179-186 Curlycup gumweed is used as an ornamental plant Since the taxonomic relationships in Grindelia genus due to the intense yellow colour of its flowers as well as have not yet been clarified completely, they are revised of their persistence over a long period of time, even in permanently (TADEY et al., 2009; DEBLE and OLIVEIRA- poor and dry soils. Because of its deep and extensive DEBLE, 2010), this process giving reasons for getting root system and of its high ability to survive and thoroughly into chromosome analysis, by both classical grow under adverse conditions, curlycup gumweed is and molecular methods. To reach a full impact, it is utilized for the rehabilitation of disturbed sites essential that the new exciting molecular findings (http://www.fs.fed.us/database/feis/plants/forb/grisqu/all. should be fully integrated with the traditional cytoge- html). The plants are unpalatable to livestock (the netical data in order to clarify the taxonomy of unpleasant taste is given by tannins, volatile oils, a bit- Grindelia (STACE, 2000). ter saponin, and resins) and they can readily absorb There are relatively few cytogenetical works on the selenium from the soil, for which reason their presence genus. This is unfortunate since both the is considered undesirable by many farmers (BARE, 1979). Grindelia chromosome numbers and the karyotype traits are In Romania, G. squarrosa is a biennial plant, with a important criteria in establishing the evolutionary pat- deep and palar root (SIRBU and OPREA, 2008, 2011). In terns as well as in plant systematics. Nothing has been the first year of vegetation, it forms a rosette, and the published on the karyomorphology of G. squarrosa. following year many branched erect stems develop; the branches carry numerous yellow flower heads. At the Considering the invasive character of G. squarrosa, its flower base there are shiny, sticky bracts, curved down- pharmacological valences and the quasi-absence of data ward (hence the name squarrosa). The leaves are alter- on its cytogenetics, the knowledge of detailed karyotype nate and oblong, tooth-edged, gland-dotted, and gummy. patterns in this species becomes a necessity, since only The fruits are pseudoachenes, without thistledown, and the diploid chromosome number is available in litera- show a marked heteromorphism (SIRBU and OPREA, ture (PINKAVA and KEIL, 1977; LANE and HARTMAN, 2011). The plant shows a high reproductive capacity in 1996). The deciphering of the genetic constitution is an Romania, counting up to 76,000 achenes per individual essential prerequisite in establishing the species struc- per year. The disc and ray achenes are morphologically ture and polymorphism, its geographical distribution, distinct in this species, and they differ in germination systematics and evolution within the genus, at inter- rate (MCDONOUGH, 1975; SIRBU et al., 2011) (Figure 1). and intra-specific levels. Figure 1. – Grindelia squarrosa (Pursh) Dunal: a, b, c–different phenophases; d. ach- enes; e. flower (photographer: SIRBU, 2011). 180 DOI:10.1515/sg-2012-0023 edited by Thünen Institute of Forest Genetics Truta et. al.·Silvae Genetica (2012) 61-4/5, 179-186 Because G. squarrosa has not been studied yet from a in the descending order of their length. Average data for karyomorphological point of view and because it is nec- each karyotype – included in tables – are the results of essary to know whether the invasive character and the the metric determinations realized on five new habitat conditions of Romania have induced some metaphases/genotype. changes at the level of the genetic material, the purpos- es of this paper are: (1) to establish the chromosome Karyotype asymmetry diploid number, (2) to detail the morphological traits of To evaluate the karyotype asymmetry, the following somatic chromosomes, (3) to construct the karyotypes, indexes were analyzed: TF %, AsI %, A1 and A2. and (4) to construct the idiogram, providing thus valu- (synonymous with ) (ARANO and able supplementary data on the chromosome constitu- AsI % index AsK % SAITO, 1980; PASZKO, 2006) represents the ratio between tion of genus. Grindelia the sum of long arm lengths of individual chromosomes and the haploid complement length: AsI %=(⌺long Material and Methods arms/haploid complement length) x 100.