Patterns of univariate and multivariate plasticity to elevated carbon dioxide in six European populations of Arabidopsis thaliana

The impact of elevated carbon dioxide on plants is a growing concern in evolutionary ecology and global change biology. Characterizing patterns of phenotypic integration and multivariate plasticity to elevated carbon dioxide can provide insights into ecological and evolutionary dynamics in future human‐altered environments. Here, we examined univariate and multivariate responses to carbon enrichment in six functional traits among six European accessions of Arabidopsis thaliana. We detected phenotypic plasticity in both univariate and multivariate phenotypes, but did not find significant variation in plasticity (genotype by environment interactions) within or among accessions. Eigenvector, eigenvalue variance, and common principal components analyses showed that elevated carbon dioxide altered patterns of trait covariance, reduced the strength of phenotypic integration, and decreased population‐level differentiation in the multivariate phenotype. Our data suggest that future carbon dioxide conditions may influence evolutionary dynamics in natural populations of A. thaliana.     

喜旱莲子草在青藏高原对模拟增温的可塑性: 引入地和原产地种群的比较

气候变暖背景下植物可通过关键性状的表型可塑性来适应环境温度的增加。表型可塑性增强进化假说预测定植到新环境中的入侵植物种群具有演化出更强表型可塑性的潜力。此前对可塑性进化的研究涵盖了外来植物性状对水分条件、光照变化、土壤养分、邻体根系以及天敌防御等的响应, 而较少有研究关注增温条件下植物重要性状的可塑性进化。已有的部分研究多集中在温带和热带地区, 而较少关注入侵植物在高寒地区对增温的响应; 且研究多集中在植物生长相关性状, 较少关注功能性状和防御性状。本研究采用同质园实验比较了喜旱莲子草6个引入地(中国)种群和6个原产地(阿根廷)种群, 在西藏拉萨模拟全天增温2℃处理下的适合度性状、功能性状和防御性状的响应差异。结果表明: (1)高寒地区模拟全天增温显著提高了喜旱莲子草总生物量(+36.4%)、地上生物量(+34.5%)、贮藏根生物量(+51.4%)和毛根生物量(+33.6%), 降低了分枝强度(-19.8%)和比茎长(-30.2%); (2)模拟全天增温使引入地种群的比叶面积和黄酮含量增加, 而原产地种群则相反。这些结果表明高寒地区全天增温2℃对喜旱莲子草可能是一种有利条件。引入地种群的适合度性状对模拟全天增温2℃的响应比原产地种群更强, 而其光能利用相关性状和防御性状的响应可能提升了其在高寒地区的适合度。因此, 在未来全球气候变暖的背景下, 高寒地区温度升高可能更有利于喜旱莲子草引入地种群的定植和扩散。     

Evolution of phenotypic plasticity a comparative approach in the phylogenetic neighbourhood of Arabidopsis thaliana

The evolution of phenotypic plasticity has rarely been examined within an explicitly phylogenetic framework, making use of modern comparative techniques. Therefore, the purpose of this study was to determine phylogenetic patterns in the evolution of phenotypic plasticity in response to vegetation shade (the ‘shade avoidance’ syndrome) in the annual plant Arabidopsis thaliana and its close relatives. Specifically, we asked the following questions: (i) Do A. thaliana and related species differ within or among clades in the magnitude and/or pattern of plasticity to shade? (ii) Are the phenotypic variance–covariance matrices (phenotypic integration) of these taxa plastic to the changes in light quality induced by the presence of a canopy? (iii) To what extent does the variation in uni- and multivariate plasticity match the phylogeny of Arabidopsis? In order to address these questions we grew individuals from six taxa of known phylogenetic relationship in a greenhouse under full sun and under a grass canopy. Taxa differed in the magnitude, but not in the pattern, of plasticities for all traits. At the univariate level, the late flowering species, A. pumila and A. griffithiana, as well as the late flowering Moscow ecotype of A. thaliana, showed greater plasticity for allocation to vegetative and reproductive meristems. At the multivariate level, several taxa displayed a very low stability of their variance–covariance structures to environmental change, with only one taxon sharing as many as three principal components across environments. We conclude that both univariate and multivariate plasticities to vegetation shade can evolve rapidly within a genus of flowering plants, with little evidence of historical constraints (phylogenetic inertia).     

Does experience with competition matter? Effects of source competitive environment on mean and plastic trait expression in Erodium cicutarium

Interspecific competition is likely to act as an agent for selection on local scales, although evidence in plants is sparse so far. We hypothesize that in annual shade-avoiding grassland species, heterogeneity in the intensity of aboveground competition for light may shape patterns of genetic variation and induce phenotypic plasticity in traits affecting competitive ability. We collected maternal seed families of Erodium cicutarium from replicated high and low competition environments and exposed them to different levels of aboveground competition in a glasshouse. We examined effects of seed source and competition treatment on expression of plant traits related to competitive ability and fitness. Source environments with high levels of competition were significantly more heterogeneous in competition intensity at both intermediate (approx. 10 m) and small (approx. 0.1 m) spatial scales. Seed source and competition treatment both had highly significant effects on trait expression. Greater intensity of competition experienced by maternal plants was coupled with lower vegetative biomass production and slower growth rates, and at the same time lower abortion rates in the offspring. We interpret these findings as an indication of greater reproductive efficiency in the next generation, in response to competition experienced by parents. There was higher total phenotypic variability in the plants from high competition source sites, but equivalent levels of phenotypic plasticity across source-site competition levels; no costs of phenotypic plasticity were detected. We concluded that differences in competition intensity can lead to trait differentiation in the next generation. For E. cicutarium, experience with competition matters: it leads to substantial phenotypic differences and more total variability in the offspring generation.     

Comparing indigenous and introduced populations of Melaleuca quinquenervia (Cav.) Blake: response of seedlings to water and pH levels

Melaleuca quinquenervia is a wetland tree species indigenous to eastern Australia. It was separately introduced to east and west Florida as an ornamental, but has since become invasive, dominating several habitat types. We tested the predictions that (1) Australian populations would exhibit more genetic variation than Florida populations, due to founder effect, and (2) high phenotypic plasticity would be found in all populations, due to the wide range of habitats occupied. We compared the phenotypic plasticity and familial variation among three Australian populations, two east Florida, and two west Florida populations in a greenhouse experiment. We grew seedlings collected from different maternal trees in each population under two water levels and three pH levels, reflecting the natural range of water levels and soil pH in Florida and Australian Melaleuca stands. We measured leaf size and shape, growth rate and above-ground biomass of seedlings and determined the components of phenotypic variance (familial, environmental, and their interaction) using univariate and multivariate analysis of variance. All traits showed significant among-population and among-family variation, as well as significant phenotypic plasticity, in response to both water level and pH level changes. Sensitivity to pH was particularly high, presumably because plants were grown under pHs ranging from 4.7 to 7.4, and because pH can influence nutrient availability. Familial variation contains genetic variation, but it may also be confounded with maternal environmental effects. Comparing Australian to Floridian Melaleuca, amounts of familial variation and phenotypic plasticity varied by trait. Overall, Australian Melaleuca had more among-population variation than Floridian Melaleuca, presumably reflecting the wider latitudinal range and longer time for evolutionary change in Australia, but had similar amounts of among-family variation, within any one population. If maternal effects are strong, among-population differences may merely reflect greater environmental differences among Australian sites than Florida sites. Australian Melaleuca had less phenotypic plasticity, possibly due to founder effects in Florida or to subsequent adaptive evolution of phenotypic plasticity in Floridian populations. Floridian Melaleuca shows little loss of familial variation, compared to indigenous Australian populations, and that, in combination with its high phenotypic plasticity, should allow it to continue to colonize new areas successfully.     

Pre-adaptation or genetic shift after introduction in the invasive species Impatiens glandulifera?

Invasive exotic plants often grow fast, reproduce rapidly and display considerable phenotypic plasticity in their invasive range, which may be essential characteristics for successful invasion. However, it remains unclear whether these characteristics are already present in native populations (pre-adaptation hypothesis) or evolve after introduction (genetic shift hypothesis).To test these hypotheses we compared means and phenotypic plasticity of vegetative and reproductive traits between populations of Impatiens glandulifera collected from either the invasive (Norway) or native range (India). Seeds were sown and the resulting plants were exposed to different experimental environments in a glasshouse. We also tested whether trait means and reaction norms harbored genetic variation, as this may promote fitness in the novel environment.We did not find evidence that invasive populations of I. glandulifera grew more vigorously or produced more seeds than native populations. Phenotypic plasticity did not differ between the native and invasive range, except for the number of nodes which was more plastic in the invasive range. Genetic variation in the slope of reaction norms was absent, suggesting that the lack of change in phenotypic plasticity between native and invasive populations resulted from low genetic variation in phenotypic plasticity initially harbored by this species. Post-introduction evolution of traits thus probably did not boost the invasiveness of I. glandulifera. Instead, the species seems to be pre-adapted for invasion.We suggest that differences in habitat between the native and invasive range, more specifically the higher nutrient availability observed in the new environment, are the main factor driving the invasion of this species. Indeed, plants in the more nutrient-rich invasive range had greater seed mass, likely conferring a competitive advantage, while seed mass also responded strongly to nutrients in the glasshouse. Interactions between habitat productivity and herbivore defense may explain the lack of more vigorous growth in the new range.     

INTER-POPULATIONAL DIFFERENTIATION AND ADAPTATION IN THE PERENNIAL,DIPLOID SPECIES FRAGARIA VESCA L.

Inter-populational differentiation and adaptation in California Fragaria vesca L. were examined using plants collected from 13 natural sites, grown in a common greenhouse and analyzed for 25 quantitative traits and 2 enzyme systems. Inter-populational variation was estimated by analysis of variance and genetic identity techniques, and micro- and macro-environmental measurements were made at each site. Stepwise multiple regressions were completed on the morphological-environmental variations and the allozyme frequency-environmental variations. Clonal propagules were compared under three controlled environmental conditions. F. vesca likely has undergone considerable ecological differentiation since significant inter-populational variation was observed in many polygenic and monogenic traits. Numerous significant correlations between environmental and character variations were found, and in many, only a few environmental variables “explained” much of that variation. Inter-populational differences in the abilities of plants to accumulate biomass and survive under salt, nutrient and shade stresses were also found. Although many of the ecotypes of F. vesca showed some phenotypic plasticity, the species is a “specialist.” Variations in root/shoot ratios occurred in biotypes grown under different conditions. Plants generally allocated more energy to aboveground biomass under shade stress and to root biomass under nutrient stress.     

Belowground mutualists and the invasive ability of Acacia longifolia in coastal dunes of Portugal

The ability to form symbiotic associations with soil microorganisms and the consequences for plant growth were studied for three woody legumes grown in five different soils of a Portuguese coastal dune system. Seedlings of the invasive Acacia longifolia and the natives Ulex europaeus and Cytisus grandiflorus were planted in the five soil types in which at least one of these species appear in the studied coastal dune system. We found significant differences between the three woody legumes in the number of nodules produced, final plant biomass and shoot ¹⁵N content. The number of nodules produced by A. longifolia was more than five times higher than the number of nodules produced by the native legumes. The obtained ¹⁵N values suggest that both A. longifolia and U. europaeus incorporated more biologically-fixed nitrogen than C. grandiflorus which is also the species with the smallest distribution. Finally, differences were also found between the three species in the allocation of biomass in the different studied soils. Acacia longifolia displayed a lower phenotypic plasticity than the two native legumes which resulted in a greater allocation to aboveground biomass in the soils with lower nutrient content. We conclude that the invasive success of A. longifolia in the studied coastal sand dune system is correlated to its capacity to nodulate profusely and to use the biologically-fixed nitrogen to enhance aboveground growth in soils with low N content.     

Different gardens, different results: native and introduced populations exhibit contrasting phenotypes across common gardens

Invasive plants may respond through adaptive evolution and/or phenotypic plasticity to new environmental conditions where they are introduced. Although many studies have focused on evolution of invaders particularly in the context of testing the evolution of increased competitive ability (EICA) hypothesis, few consistent patterns have emerged. Many tests of the EICA hypothesis have been performed in only one environment; such assessments may be misleading if plants that perform one way at a particular site respond differently across sites. Single common garden tests ignore the potential for important contributions of both genetic and environmental factors to affect plant phenotype. Using a widespread invader in North America, Cynoglossum officinale, we established reciprocal common gardens in the native range (Europe) and introduced range (North America) to assess genetically based differences in size, fecundity, flowering phenology and threshold flowering size between native and introduced genotypes as well as the magnitude of plasticity in these traits. In addition, we grew plants at three nutrient levels in a pot experiment in one garden to test for plasticity across a different set of conditions. We did not find significant genetically based differences between native and introduced populations in the traits we measured; in our experiments, introduced populations of C. officinale were larger and more fecund, but only in common garden experiments in the native range. We found substantial population-level plasticity for size, fecundity and date of first flowering, with plants performing better in a garden in Germany than in Montana. Differentiation of native populations in the magnitude of plasticity was much stronger than that of introduced populations, suggesting an important role for founder effects. We did not detect evidence of an evolutionary change in threshold flowering size. Our study demonstrates that detecting genetically based differences in traits may require measuring plant responses to more than one environment.     

本地和外来草本物种对水分条件时间异质性的可塑响应

极端气候导致的干旱和水淹事件频发，影响了外来植物和本地植物的生长。为了解外来种和本地种植物对干旱和水淹事件发生顺序的响应，探讨草本植物适应水分时间异质性的策略，该文以美国蒙大拿州西部4种本地植物和4种外来植物为研究对象，将所有植物分别进行持续湿润(对照，CK)、水淹-干旱(I-D)和干旱-水淹(D-I)处理，并观测一系列形态和生物量特征的变化。结果表明：(1)与CK相比，D-I和I-D处理均显著降低了外来种的总生物量(P<0.05)。(2)D-I显著降低了本地种早期总生物量、后期地下生物量和根冠比，但显著提高了其后期的相对生长(P<0.05)。(3)D-I处理显著降低了所有植物的地下-地上生物量关系的异速指数，外来种异速指数显著高于本地种(P<0.05)。综上认为，极端事件(水淹和干旱)的发生顺序能改变外来植物和本地植物的生物量分配，早期干旱比后期干旱更容易减少植物生物量的积累，但能促进本地种后期的生长；本地种在环境胁迫下不被降低的总生物量表现说明维持表型稳定的能力较强；D-I处理下本地种和外来种地上和地下生物量关系的分配方式不同。     

Ecophysiological basis of the Jack-and-Master strategy: Taraxacum officinale (dandelion) as an example of a successful invader

Aims Successful invasive plants are often assumed to display significant levels of phenotypic plasticity. Three possible strategies by which phenotypic plasticity may allow invasive plant species to thrive in changing environments have been suggested: (i) via plasticity in morphological or physiological traits, invasive plants are able to maintain a higher fitness than native plants in a range of environments, including stressful or low-resource habitats: a 'Jack-of-all-trades' strategy; (ii) phenotypic plasticity allows the invader to better exploit resources available in low stress or favorable habitats, showing higher fitness than native ones: a 'Master-of-some' strategy and (iii) a combination of these abilities, the 'Jack-and-Master' strategy.Methods We evaluated these strategies in the successful invader Taraxacum officinale in a controlled experiment mimicking natural environmental gradients. We set up three environmental gradients consisting of factorial arrays of two levels of temperature/light, temperature/water and light/water, respectively. We compared several ecophysiological traits, as well as the reaction norm in fitness-related traits, in both T. officinale and the closely related native Hypochaeris thrincioides subjected to these environmental scenarios.Important findings Overall, T. officinale showed significantly greater accumulation of biomass and higher survival than the native H. thrincioides, with this difference being more pronounced toward both ends of each gradient. T. officinale also showed significantly higher plasticity than its native counterpart in several ecophysiological traits. Therefore, T. officinale exhibits a Jack-and-Master strategy as it is able to maintain higher biomass and survival in unfavorable conditions, as well as to increase fitness when conditions are favorable. We suggest that this strategy is partly based on ecophysiological responses to the environment, and that it may contribute to explaining the successful invasion of T. officinale across different habitats.     

喜旱莲子草对模拟全天增温的可塑性: 引入地和原产地种群的比较

植物可以通过关键功能性状的表型可塑性来适应气候变暖背景下环境温度的增加。表型可塑性增强进化假说(evolution of increased phenotypic plasticity hypothesis)认为外来植物在引入地进化出了更强的表型可塑性。以往对该假说的验证多集中于外来植物对光照、水分、养分、邻体以及天敌等的可塑性进化, 而对增温条件下植物生长和功能性状可塑性进化的研究相对较少。仅有的几项研究多集中在温带地区, 且多集中于研究植物生长相关的性状, 而对植物的抗性和草食作用对增温的响应的关注相对较少。本研究采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)引入地(中国)和原产地(阿根廷)各8个种群的生物量、功能性状和草食作用在热带地区(广州市增城区)对模拟全天增温2℃的响应差异。结果表明: (1)模拟全天增温显著降低了喜旱莲子草总生物量(-7.8%)、贮藏根生物量(-12.8%)、分枝强度(-11.6%)和茎端取食率(-34.4%)。(2)模拟全天增温造成的引入地种群总生物量降低幅度大于原产地种群; 模拟全天增温使引入地种群的比茎长和茎端取食率降低, 而原产地种群则相反。(3)无论是否模拟全天增温, 引入地种群的贮藏根生物量(+31.5%)、分枝强度(+38.5%)、比茎长(+30.2%)、根冠比(+24.5%)和比叶面积(+20.0%)均高于原产地种群, 茎端取食率则低于原产地种群(-35.8%)。这些结果表明, 热带地区全天增温2℃对喜旱莲子草是一种胁迫; 引入地种群的生物量对模拟全天增温2℃的响应更强, 而其株形相关性状(比茎长)和草食作用(茎端取食率)对模拟全天增温的可塑性方向与原产地种群相反。由于引入地种群在热带地区模拟全天增温条件下生物量的下降和草食作用的增加明显高于原产地种群, 因此在未来全球气候变暖的背景下, 热带地区温度升高可能不利于喜旱莲子草种群多度的增加。     

两种入侵植物与三种本地植物根系特征的比较研究

以菊科2种入侵植物飞机草(Chromolaena odorata (L.)R. M. King & H. Rob)和紫茎泽兰( Eupatorium adenophorum Speng)以及生活型相似的3种本地植物异叶泽兰(Eupatorium heterophyllum DC.)、佩兰(Eupa-torium for...     

The effect of steepness of temporal resource gradients on spatial root allocation

Plants are able to discriminately allocate greater biomass to organs that grow under higher resource levels. Recent evidence demonstrates that split-root plants also discriminately allocate more resources to roots that grow under dynamically improving nutrient levels, even when their other roots grow in richer patches. Here, we further tested whether, besides their responsiveness to the direction of resource gradients, plants are also sensitive to the steepness of environmental trajectories. Split-root Pisum sativum plants were grown so that one of their roots developed under constantly-high nutrient levels and the other root was subjected to dynamically improving nutrient levels of variable steepness. As expected, plants usually allocated a greater proportion of their biomass to roots that developed under constantly high resource availability; however, when given a choice, they allocated greater biomass to roots that initially experienced relatively low but steeply improving nutrient availabilities than to roots that developed under continuously-high nutrient availability. Such discrimination was not observed when the roots in the poor patch experienced only gentler improvements in nutrient availability. The results are compatible with the notion that responsiveness to the direction and steepness of environmental gradients could assist annual plants to increase their performance by anticipating resource availabilities foreseeable before the end of the growing season. The results exemplify the ability of plants to integrate and utilize environmental information and execute adaptive behaviors which, until recently, were attributed only to animals with central nervous systems.Key words: anticipatory responses, environmental information, foraging, gradient perception, phenotypic plasticity, plant behavior, rate of change, resource gradients     

Nutrient enrichment enhances hidden differences in phenotype to drive a cryptic plant invasion

Many mechanisms of invasive species success have been elucidated, but those driving cryptic invasions of non‐native genotypes remain least understood. In one of the most successful cryptic plant invasions in North America, we investigate the mechanisms underlying the displacement of native Phragmites australis by its Eurasian counterpart. Since invasive Phragmites’ populations have been especially prolific along eutrophic shorelines, we conducted a two‐year field experiment involving native and invasive genotypes that manipulated nutrient level and competitor identity (inter‐ and intra‐genotypic competition) to assess their relative importance in driving the loss of native Phragmites. Inter‐genotypic competition suppressed aboveground biomass of both native and invasive plants regardless of nutrient treatment (～ 27%), while nutrient addition disproportionately enhanced the aboveground biomass (by 67%) and lateral expansion (by > 3 × farther) of invasive Phragmites. Excavation of experimental plots indicated that nutrient addition generates these differences in aboveground growth by differentially affecting rhizome production in invasive vs native plants; invasive rhizome biomass and rhizome length increased by 595% and 32% with nutrient addition, respectively, while natives increased by only 278% and 15%. Regardless of nutrient level, native rhizomes produced twice as many roots compared to invasives, which field surveys revealed are heavily infected with mycorrhizal symbionts. These results suggest that native Phragmites competes well under nutrient‐limited conditions because its rhizomes are laden with nutrient‐harvesting roots and mycorrhizae. Invasive Phragmites’ vigorous aboveground response to nutrients and scarcity of lateral roots, in contrast, may reflect its historic distribution in eutrophic Eurasian wetlands and correspond to its prevalence in New England marshes characterized by elevated nutrient availability and relaxed nutrient competition. These findings reveal that discrete differences in phenotype can interact with anthropogenic modification of environmental conditions to help explain the success of cryptic invaders.     

Environmental Heterogeneity and Population Differentiation in Plasticity to Drought in <Emphasis Type="Italic">Convolvulus Chilensis</Emphasis> (Convolvulaceae)

Plant populations may show differentiation in phenotypic plasticity, and theory predicts that greater levels of environmental heterogeneity should select for higher magnitudes of phenotypic plasticity. We evaluated phenotypic responses to reduced soil moisture in plants of Convolvulus chilensis grown in a greenhouse from seeds collected in three natural populations that differ in environmental heterogeneity (precipitation regime). Among several morphological and ecophysiological traits evaluated, only four traits showed differentiation among populations in plasticity to soil moisture: leaf area, leaf shape, leaf area ratio (LAR), and foliar trichome density. In all of these traits plasticity to drought was greatest in plants from the population with the highest interannual variation in precipitation. We further tested the adaptive nature of these plastic responses by evaluating the relationship between phenotypic traits and total biomass, as a proxy for plant fitness, in the low water environment. Foliar trichome density appears to be the only trait that shows adaptive patterns of plasticity to drought. Plants from populations showing plasticity had higher trichome density when growing in soils with reduced moisture, and foliar trichome density was positively associated with total biomass. Co-ordinating editor: F. Stuefer     

Traits correlate with invasive success more than plasticity: A comparison of three Centaurea congeners

The importance of phenotypic plasticity for successful invasion by exotic plant species has been well studied, but with contradictory and inconclusive results. However, many previous studies focused on comparisons of native and invasive species that co‐occur in a single invaded region, and thus on species with potentially very different evolutionary histories. We took a different approach by comparing three closely related Centaurea species: the highly invasive C. solstitialis, and the noninvasive but exotic C. calcitrapa and C. sulphurea. These species have overlapping distributions both in their native range of Spain and in their non‐native range of California. We collected seeds from 3 to 10 populations from each region and species and grew them in common garden greenhouse conditions to obtain an F1 generation in order to reduce maternal effects. Then, F1 seeds were grown subjected to simulated herbivory, variation in nutrient availability, and competition, to explore plasticity in the responses to these conditions. We found little variation in phenotypic plasticity among species and regions, but C. solstitialis plants from California produced more biomass in competition than their Spanish conspecifics. This species also had the highest relative growth rates when in competition and when grown under low nutrient availability. Noninvasive congeners produced intermediate or opposite patterns.     

Consequences of phenotypic plasticity vs. interspecific differences in leaf and root traits for acquisition of aboveground and belowground resources

Trade-offs between acquisition capacities for aboveground and belowground resources were investigated by studying the phenotypic plasticity of leaf and root traits in response to different irradiance levels at low nutrient supply. Two congeneric grasses with contrasting light requirements, Dactylis glomerata and D. polygama, were used. The aim was to analyze phenotypic covariation in components of leaf area and root length in response to above- and belowground resource limitation and the consequences of this variation for resource acquisition and plant growth. At intermediate shading (30 and 20% of full sunlight) the plants were able to maintain their total root length, despite a strongly increased total leaf area and a reduced biomass allocation to roots. This was associated with an unaltered or slightly increased nutrient uptake and growth. At 5.5% relative irradiance, growth was severely reduced, especially in the shade-tolerant D. polygama. The results show that constraints on acquisition capacities for aboveground and belowground resources, caused by biomass allocation, may be alleviated by plasticity in other traits such as tissue-mass density and thickness of roots and leaves. The results also suggest different adaptive constraints for phenotypic plasticity and for genetically determined interspecific variation. Phenotypic plasticity tends to maximize resource acquisition and growth rate in the short term, whereas the higher tissue-mass density and the longer leaf life-span of shade-tolerant species indicate reduced loss rates as a more advantageous species-specific adaptation to shade in the long term.     

Phenotypic plasticity as a clue for invasion success of the submerged aquatic plant Elodea nuttallii

• Two closely related alien submerged aquatic plants were introduced into Europe. The new invader (Elodea nuttallii) gradually displaced E. canadensis even at sites where the latter was well established. The aim of the study was to evaluate the combined effects of environmental factors on several phenotypic characteristics of the two Elodea species, and to relate these phenotypic characteristics to the invasion success of E. nuttallii over E. canadensis.
• In a factorial design, Elodea plants were grown in aquaria containing five different nitrogen concentrations and incubated at five different light intensities. We used six functional traits (apical shoot RGR), total shoot RGR, relative elongation, root length, lateral spread, branching degree) to measure the environmental response of the species. We calculated plasticity indices to express the phenotypic differences between species.
• Light and nitrogen jointly triggered the development of phenotypic characteristics that make E. nuttallii a more successful invader in eutrophic waters than E. canadensis. The stronger invader showed a wider range of phenotypic plasticity. The apical elongation was the main difference between the two species, with E. nuttallii being more than two times longer than E. canadensis. E. canadensis formed dense side shoots even under high shade and low nitrogen levels, whereas E. nuttallii required higher light and nitrogen levels.
• We found that under more eutrophic conditions, E. nuttallii reach the water surface sooner than E. canadensis and through intensive branching outcompetes all other plants including E. canadensis. Our findings support the theory that more successful invaders have wider phenotypic plasticity.

     

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.