Ecophysiological plasticity and local differentiation help explain the invasion success of Taraxacum officinale (dandelion) in South America

Plasticity and local adaptation have been suggested as two main mechanisms that alien species use to successfully tolerate and invade broad geographic areas. In the present study, we try answer the question if the mechanism for the broad distributional range of T. officinale is for phenotypic plasticity, ecotypic adaptation or both. For this, we used individuals of T. officinale originated from seeds collected in five localities along its latitudinal distribution range in the southern‐hemisphere. Seedlings were acclimated at 5 and 25°C for one month. After the acclimation period we evaluated ecophysiological and cytogenetic traits. Additionally, we assessed the fitness at each temperature by recording the seed output of individuals from different localities. Finally, we performed a manipulative experiment in order to assess the tolerance to herbivory and competitive ability between T. officinale from all origins and Hypochaeris scorzonerae a co‐occurring native species. Overall, individuals of T. officinale showed high plasticity and ecotypic adaptation for all traits assessed in this study. Changes both in physiology and morphology observed in T. officinale from different origins were mostly correlated, enhancing their ecophysiological performance in temperatures similar to those of their origin. Additionally, all localities showed the same chromosome number and ploidy level. On the other hand, all individuals showed an increase the seed output at 25°C, but those from northern localities increased more. T. officinale from all origins was not significantly affected by herbivory while native showed a negative effect. On the other hand, T. officinale exerted a strong negative effect on the native species, but this former not effected significantly to the invasive T. officinale. High plasticity and local adaptation in all ecophysiological traits, seed‐set and the low cytogenetic variability in T. officinale suggests that both strategies are present in this invasive plant species and are not mutually exclusive. Finally, higher tolerance to herbivory and competitive ability suggests that T. officinale could perform successfully in environments with different climatic conditions, and thus colonize and invade South‐America.     

Detecting small-scale genotype-environment interactions in apomictic dandelion (Taraxacum officinale) populations

Studies of genotype × environment interactions (G × E) and local adaptation provide critical tests of natural selection’s ability to counter opposing forces such as gene flow. Such studies may be greatly facilitated in asexual species, given the possibility for experimental replication at the level of true genotypes (rather than populations) and the possibility of using molecular markers to assess genotype–environment associations in the field (neither of which is possible for most sexual species). Here, we tested for G × E in asexual dandelions (Taraxacum officinale) by subjecting six genotypes to experimental drought, mown and benign (control) conditions and subsequently using microsatellites to assess genotype–environment associations in the field. We found strong G × E, with genotypes that performed poorly under benign conditions showing the highest performance under stressful conditions (drought or mown). Our six focal genotypes comprise > 80% of plants in local populations. The most common genotype in the field showed its highest relative performance under mown conditions (the most common habitat in our study area), and almost all plants of this genotype in the field were found growing in mowed lawns. Genotypes performing best under benign experimental conditions were found most frequently in unmown conditions in the field. These results are strongly indicative of local adaptation at a very small scale, with unmown microsites of only a few square metres typically embedded within larger mown lawns. By studying an asexual species, we were able to map genotypes with known ecological characteristics to environments with high spatial precision.     

Sclerotinia minor avances fruiting and reduces germination in dandelion (Taraxacum officinale)

Sclerotinia minor Jagger is a promising biocontrol agent for dandelion in turfgrass. When a flowering dandelion population was treated with S. minor, flowering accelerated to the fruiting stage within 4 days. This developmental response was 4-5 days earlier than in the control, untreated plants and was not observed in herbicide-treated plants. Seeds obtained from the fungal-treated plants were smaller, lighter and their germination rate was reduced by 48.4 and 47.3% for spring and fall applications, respectively. S. minor was not detected in dandelion seeds from the fungal-treated plants. In addition to effective control of mature (flowering) dandelions, seeds dispersed by dying plants have reduced germination and are not transferring S. minor off target.     

Development and characterization of microsatellite markers in the sexual-apomictic complex Taraxacum officinale (dandelion)

 Microsatellite markers were developed in Taraxacum officinale to study gene flow between sexual and apomictic plants and to identify clones. Twenty five thousand genomic DNA clones were hybridized with a (CT)₁₂D probe. The density of (GA/CT) _n repeats was estimated at one every 61 kb in the T. officinale genome, which translates to 13 500 repeats per haploid genome. Ninety two percent of 110 positive clones sequenced contained at least one (GA/CT) _n≥5 repeat. Sixteen (CA/GT) _n≥5 and 11 (AT) _n≥5 arrays were also found in these sequences, suggesting some clustering of dinucleotide repeats. Among 50 PCR primer pairs tested, 32 produced bands and 28 of them were polymorphic. Of these polymorphic markers, 15 were putatively single-locus and the other 13 produced only polymorphic fingerprints. Six loci were further characterized for polymorphism and showed between 6 and 32 alleles per locus. Among eight primer pairs used to analyze the progeny of a sexual cross, seven were co-dominant single-locus Mendelian markers, but one (MSTA10) gave a dominant pattern in accordance with the hypothesis of a null allele segregating in a Mendelian fashion. Three pairs of loci among 28 showed significant linkages of 10, 21, and 39 cM. Observed and expected heterozygosities in two sexual populations indicate that null alleles may be present at two loci, including MSTA10. Received: 4 November 1997 / Accepted: 12 February 1998     

Clonal variation in floral stage timing in the common dandelion Taraxacum officinale (Asteraceae)

We investigated the hypothesis that dandelion clones (Taraxacum officinale Weber, sensu lato; Asteraceae) differ in their floral stage timing characteristics under a constant set of environmental conditions. To test this hypothesis, plants representing nine different dandelion clones (identified by DNA fingerprinting) were grown in groups of five (N = 45) in a growth chamber for a period of 8 mo, with chamber settings similar to environmental conditions at peak dandelion flowering time for their population sites. Five flowering phenology parameters were monitored daily for a total of 301 buds developing during this time: (1) time to bud; (2) time to full opening and inflorescence maturation (i.e., first anthesis); (3) time to re-closure of an inflorescence; (4) time to fruit (full re-opening of the inflorescence); and (5) total flowering time. Scape length at the appearance of a fully expanded infructescence was also measured for each individual. Significant differences in mean time to inflorescence, mean time to re-closure, mean time to fruit, and mean total flowering time were revealed among some dandelion clones (Kruskal-Wallis, P ≤ 0.0005). No differences in mean number of inflorescence buds per plant (P = 0.2217), mean time to bud (P = 0.2396), or mean scape length (P = 0.3688) were detected among the nine clones. These results suggest that differences in floral stage timing may in part involve varying genotypic environmental response characteristics and that these differences may have potential fitness effects. Further research is needed to determine if such clonal differences are observed under a broader range of uniform environmental conditions.     

A comparison of phenotypic plasticity in the native dandelion Taraxacum ceratophorum and its invasive congener T. officinale

We compared plastic responses to variation in the light environment for sympatric populations of native and exotic dandelion species, Taraxacum ceratophorum and Taraxacum officinale. Plasticity in leaf size, inflorescence height, reproductive phenology and dispersal-related traits were measured under experimentally altered light quality (red : far-red light ratio, R : FR) and light intensity (photosynthetically active radiation, PAR). To test whether differences in means and reaction norms of dispersal-related traits between species affected colonization potential, we created seed-dispersal models based on seed-fall rate and release height. Differences in plasticity between species were not systematic, but varied in direction and magnitude among traits. Taraxacum officinale produced larger leaves that exhibited greater plasticity in size under variable light intensity than T. ceratophorum. Plasticity in scape length at flowering occurred in relation to R : FR ratio in both species, but tended to be greater in T. ceratophorum. Seed-bearing scapes of T. officinale were taller and more canalized in height across light regimes than scapes of T. ceratophorum. Seeds of T. officinale were smaller than seeds of T. ceratophorum. Models predict greater dispersal in T. officinale within open and vegetated habitats. In contrast to the idea that plasticity promotes invasiveness, results suggest that the lack of plasticity in dispersal-related traits enhances the colonization potential of T. officinale.     

Photonastic and thermonastic opening of capitulum in dandelion,Taraxacum officinale andTaraxacum japonicum

The capitula ofTaraxacum officinale andT. japonicum open in response to temperature rise at lower temperatures (thermonasty), and in response to light at higher temperatures (photonasty), as was the case inT. albidum. The capitula ofT. officinale could respond to the same temperature rise more sensitively than those ofT. albidum orT. japonicum. The minimum temperature for photonastic opening is as low as 13 C forT. officinale, while that forT. albidum andT. japonicum is about 18 C. That is why the capitula ofT. officinale opened earlier than those ofT. albidum andT. japonicum in the morning in April under natural conditions. The capitulum continued to be open for about 13–14 hr inT. officinale and about 8–11 hr inT. japonicum and inT. albidum both under natural conditions in April and even under constant light-temperature conditions, suggesting that the time of capitula-closing in these three species is not controlled by changes in environmental factors (light and/or temperature).     

Drought tolerance in the alpine dandelion, Taraxacum ceratophorum (Asteraceae), its exotic congener T. officinale, and interspecific hybrids under natural and experimental conditions

We compared water relations and adaptations to drought stress in native and invasive exotic dandelions, Taraxacum ceratophorum and T. officinale. Photosynthesis (A), transpiration (E), and water use efficiency (WUE; carbon gained/water lost) were measured for the two species under extreme drought in the alpine tundra of Colorado, USA. We also subjected both species and F(1) hybrids to a dry-down experiment to determine how relative physiological performance varied with water availability. Photosynthesis and transpiration in the field were low and did not differ between Taraxacum congeners; however, native T. ceratophorum had higher WUE than T. officinale. After 6 days of greenhouse drought, photosynthesis and transpiration were reduced in T. officinale compared to T. ceratophorum. Taraxacum ceratophorum maintained high WUE under control and drought treatments. Conversely, WUE in T. officinale was highly plastic between watered (low WUE) and dry-down (high WUE) treatments. Hybrids did not exhibit heterosis; instead, they were similar to T. officinale in A and E and intermediate to the parental species in WUE. Overall, results suggest that native dandelions are more drought tolerant than invasive congeners or their hybrids, but have less plasticity in WUE. Arid habitats and occasional drought in mesic sites may provide native dandelions with refugia from negative interactions with invasives.     

Statistical downscaling of general-circulation-model- simulated average monthly air temperature to the beginning of flowering of the dandelion (Taraxacum officinale) in Slovenia

Phenological observations are a valuable source of information for investigating the relationship between climate variation and plant development. Potential climate change in the future will shift the occurrence of phenological phases. Information about future climate conditions is needed in order to estimate this shift. General circulation models (GCM) provide the best information about future climate change. They are able to simulate reliably the most important mean features on a large scale, but they fail on a regional scale because of their low spatial resolution. A common approach to bridging the scale gap is statistical downscaling, which was used to relate the beginning of flowering of Taraxacum officinale in Slovenia with the monthly mean near-surface air temperature for January, February and March in Central Europe. Statistical models were developed and tested with NCAR/NCEP Reanalysis predictor data and EARS predict and data for the period 1960-1999. Prior to developing statistical models, empirical orthogonal function (EOF) analysis was employed on the predictor data. Multiple linear regression was used to relate the beginning of flowering with expansion coefficients of the first three EOF for the Janauary, Febrauary and March air temperatures, and a strong correlation was found between them. Developed statistical models were employed on the results of two GCM (HadCM3 and ECHAM4/OPYC3) to estimate the potential shifts in the beginning of flowering for the periods 1990-2019 and 2020-2049 in comparison with the period 1960-1989. The HadCM3 model predicts, on average, 4 days earlier occurrence and ECHAM4/OPYC3 5 days earlier occurrence of flowering in the period 1990-2019. The analogous results for the period 2020-2049 are a 10- and 11-day earlier occurrence.     

A new subtilisin-like proteinase from roots of the dandelion Taraxacum officinale Webb S. L.

A serine proteinase from roots of Taraxacum officinale Webb S. L. was isolated by affinity chromatography and gel-filtration on Superose 6R using FPLC. The enzyme is a 67-kD glycoprotein containing 54% carbohydrate which we have named taraxalisin. The substrate specificity of taraxalisin toward synthetic peptides and oxidized insulin B-chain is comparable with that of cucumisin from Cucumis melo and the subtilisin-like serine proteinase macluralisin from Maclura pomifera. The proteinase is inactivated by DFP and PMSF. Taraxalisin exhibits maximal activity at pH 8.0. The pH range for stability of the enzyme is narrow--6.0-9.0. The temperature optimum for the subtilisin-like activity is 40 degrees C. The N-terminal sequence of taraxalisin has 40% of its residues identical to those of subtilisin Carlsberg. Thus, the serine proteinase from dandelion roots is a member of the subtilisin family, which is evidently widespread in the plant kingdom.     

Discovery of novel antimicrobial peptides with unusual cysteine motifs in dandelion Taraxacum officinale Wigg. flowers

Three novel antimicrobial peptides designated ToAMP1, ToAMP2 and ToAMP3 were purified from Taraxacum officinale flowers. Their amino acid sequences were determined. The peptides are cationic and cysteine-rich and consist of 38, 44 and 42 amino acid residues for ToAMP1, ToAMP2 and ToAMP3, respectively. Importantly, according to cysteine motifs, the peptides are representatives of two novel previously unknown families of plant antimicrobial peptides. ToAMP1 and ToAMP2 share high sequence identity and belong to 6-Cys-containing antimicrobial peptides, while ToAMP3 is a member of a distinct 8-Cys family. The peptides were shown to display high antimicrobial activity both against fungal and bacterial pathogens, and therefore represent new promising molecules for biotechnological and medicinal applications.     

Phenotypic plasticity and performance of <Emphasis Type="Italic">Taraxacum officinale</Emphasis> (dandelion) in habitats of contrasting environmental heterogeneity

Ecological theory predicts a positive association between environmental heterogeneity of a given habitat and the magnitude of phenotypic plasticity exhibited by resident plant populations. Taraxacum officinale (dandelion) is a perennial herb from Europe that has spread worldwide and can be found growing in a wide variety of habitats. We tested whether T. officinale plants from a heterogeneous environment in terms of water availability show greater phenotypic plasticity and better performance in response to experimental water shortage than plants from a less variable environment. This was tested at both low and moderate temperatures in plants from two sites (Corvallis, Oregon, USA, and El Blanco, Balmaceda, Chile) that differ in their pattern of monthly variation in rainfall during the growth season. We compared chlorophyll fluorescence (photosynthetic performance), flowering time, seed output, and total biomass. Plants subjected to drought showed delayed flowering and lower photosynthetic performance. Plants from USA, where rainfall variation during the growth season was greater, exhibited greater plasticity to water shortage in photosynthetic performance and flowering time than plants from Chile. This was true at both low and moderate temperatures, which were similar to early- and late-season conditions, respectively. However, phenotypic plasticity to decreased water availability was seemingly maladaptive because under both experimental temperatures USA plants consistently performed worse than Chile plants in the low water environment, showing lower total biomass and fewer seeds per flower head. We discuss the reliability of environmental clues for plasticity to be adaptive. Further research in the study species should include other plant traits involved in functional responses to drought or potentially associated with invasiveness.     

Constitutive knox1 gene expression in dandelion (Taraxacum officinale, Web.) changes leaf morphology from simple to compound

Seed plants with compound leaves constitute a polyphyletic group, but studies of diverse taxa show that genes of the class 1 KNOTTED-LIKE HOMEOBOX (KNOX1) family are often involved in compound leaf development. This suggests that knox1 genes have been recruited on multiple occasions during angiosperm evolution (Bharathan et al. in Science 296:1858–1860, 2002). In agreement with this, we demonstrate that the simple leaf of dandelion (Taraxacum officinale Web.) can be converted into a compound leaf by the constitutive expression of heterologous knox1 genes. Dandelion is a rosette plant of the family Asteraceae, characterised by simple leaves with deeply lobed margins and endogenous knox1 gene expression. Transgenic dandelion plants constitutively expressing the barley (Hordeum vulgare L.) hooded gene (bkn3, barley knox3) or the related bkn1 gene, developed compound leaves featuring epiphyllous rosettes. We discuss these results in the context of two current models of compound leaf formation.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Enhanced fitness and greater herbivore resistance: implications for dandelion invasion in an alpine habitat

Several hypotheses have been proposed to explain the defense strategies of invasive plants in new ranges. In the absence of specialist herbivores, it is believed that invasive plants may allocate fewer resources to resistance and more to growth and reproduction, thus increasing tolerance to damage in the invasive genotypes. In order to test these predictions, we compared both performance (growth and reproduction) and defense strategies (tolerance and resistance) of two populations of Taraxacum officinale, one from the native range in the French Alps, and one from the introduced range in the Chilean Andes. Individuals from the introduced population demonstrated improved reproductive traits relative to those from the native population, although there was no discernible difference in biomass accumulation. Additionally, reduced tolerance was evident in the case of the former; whereas fitness traits of native plants were unaffected by damage, invasive plants reduced growth and seed output by 25 and 30% respectively following damage treatments. Increases in levels of phenols and anthocyanins, produced as a defense response to herbivory, were observed in introduced plants. Our results suggest that reallocation of resources to reproduction may be an important factor favouring invasive success of T. officinale in Chile, and that a higher investment in chemical resistance traits in this population may also be a factor in this regard.     

The potential for genetic assimilation of a native dandelion species, Taraxacum ceratophorum (Asteraceae), by the exotic congener T. officinale

Exotic plant species can threaten closely related native congeners through asymmetric hybridization and subsequent backcrossing, the process known as genetic assimilation. I explore the initial stages of this process in Taraxacum ceratophorum (Asteraceae), the native alpine dandelion, and the invasive apomict T. officinale. In central Colorado, seven T. ceratophorum populations all occur in sympatry with T. officinale. In one large population on Pennsylvania Mountain, surveys further revealed that flowering phenologies and visiting insect taxa overlap almost completely for both Taraxacum species. Together these results indicated that heterospecific pollen transfer is likely. Crossing experiments showed that T. ceratophorum is an obligate outcrosser, and interspecific hand pollinations resulted in 37.3% seed set. However, molecular analysis of the F1 offspring indicated that only 33.2% of germinating seeds were hybrids; the remainder were selfed offspring produced from a breakdown in self-incompatibility (the mentor effect). Although the mentor effect helps reduce the production of hybrids, the asymmetrical direction of hybridization creates the potential for genetic assimilation of T. ceratophorum by T. officinale.     

Phenotypic plasticity in alpine Erigeron species (Asteraceae)

Utelli, A.-B., Huber, W. & Zopfi, H.-J. 1995. Phenotypic plasticity in alpine Erigeron species (Asteraceae). — Nord. J. Bot. 15: 483–492. Copenhagen. ISSN 0107–055X.
Twelve morphological characters were analysed in 67 E. alpinus, E. neglectus and E. uniflorus plants which had been transplanted from their natural habitat to the greenhouse. Mean values and standard deviations were compared for each species. The phenotypic plasticity of the characters resp. their genetic basis was estimated from the correlation of morphological data recorded on single individuals, each grown under both ecological conditions.
A considerable phenotypic plasticity was found in all species and for most characters. E. unijlorus was the most constant species regarding taxonomically relevant characters such as pubescence of the basal leaves, number of flowering heads and number of filiform flowers. In all three species the same highly significant directional variation in plant height and involucre length was found which suggests that these characters have some genetic basis in Erigeron .