Phenotypic Plasticity in the Annual Weed Polygonum aviculare

Reaction norms of fourteen life history and morphological traits were investigated in four tetra- and two hexaploid genotypes of the annual weed species complex, Polygonum aviculare. The plants were cultivated in six treatments consisting of factorial combinations of three pot sizes and two fertility levels. All characters, except life span, were plastic but the relative importance of genotype (G), treatment (T) and interaction (G × T) to total variance was strongly trait-specific. Consistent genetic differentiation, not correlated with ploidy level, was found in metamer size and life history: genotypes originating from trampled sites had smaller metamers and shorter shoots while those originating from sites with a short growing season, due to weeding activities, had a shorter life span, an earlier flowering date and a higher biomass allocation to reproduction compared to genotypes from less disturbed sites. Significant variation was found in reaction norms for all characters, including a lower amount of plasticity in metamer size in genotypes with numerous metamers and a lower amount of plasticity in total weight in shortlived genotypes. This suggested that variation in phenotypic plasticity reflected developmental constraints imposed by contrasting life span and metamer size in different genotypes. There was no evidence for niche differentiation along the soil resource gradient, suggesting that the species is comprised of “general purpose” genotypes with respect to soil fertility. It is concluded that the Polygonum aviculare complex has evolved a “dual” adaptive strategy i.e. a combination of genetic polymorphism and high phenotypic plasticity.     

Variation of development time until flowering in natural populations ofCapsella bursa-pastoris (Cruciferae)

Seed samples were collected from wild populations ofCapsella bursa-pastoris along a transsect from Northern to Southern Europe. Progeny was grown in (a) open-field random block experiments (47 populations) and (b) in growth chambers under five to seven controlled temperature regimes (18 populations). Beginning of flowering was recorded, and great differences between and also within populations are documented. Some populations are extremely heterogenous whereas others are homogenous in this respect. Some biotypes react positively when exposed to lower temperatures, others are inhibited. In many cases specific effects of day- and/or night-temperatures can be inferred. In some progenies begin of flowering is independent of temperature as long as this exceeds the 5:10°C regimen. Altogether,Capsella bursa-pastoris displays definite intraspecific variation in time required until flowering. Adaptations to local ecological conditions are obvious. In addition to a genotypic component pronounced environmental interactions provide the plants with a component of phenotypic plasticity. The degree of modificability apparently varies itself and seems to be controlled by selection; the phenotypic plasticity, therefore, displays adaptive variation patterns, too.Adaptation in Life History Traits of Colonizing Plant Species; Part of a doctoral thesis by the first author.     

Genetic differences and phenotypic plasticity in body size between high- and low-altitude populations of the ground beetle Carabus tosanus

The body size of a univoltine carabid beetle Carabus tosanus on Shikoku Island, Japan, was clearly smaller in higher‐altitude populations (subspecies), which possibly represents incipient speciation. To explore the determinants of altitudinal differences in body size in this species, we studied the degree of phenotypic plasticity by conducting rearing experiments at two constant temperatures and examined genetic differences through interpopulation crosses. At 15 °C, C. tosanus had a longer developmental period and a shorter adult body than at 20 °C. Nevertheless, variation in body size due to temperature effects (phenotypic plasticity) was small compared to the interpopulation differences, which suggests substantial genetic differences between populations (subspecies) at different altitudes. In F₁ offspring from crosses between a low‐altitude (subspecies tosanus) and a high‐altitude population (subspecies ishizuchianus), adult body length was affected by the genotypes of both parents, with an interaction effect of parental genotype and offspring sex. Further analyses revealed that adult body length was affected by sex‐linked factors in addition to autosomal factors. These genetic differences in body size may have resulted from adaptations to different altitudes and may be important for the process of incipient speciation because body size differences could contribute to premating reproductive isolation.     

Patterns of variation of a common fern (Athyrium filix-femina; Woodsiaceae): population structure along and between altitudinal gradients

Genetic variability of Athyrium filix-femina populations was evaluated with regard to phenotypic, allozyme, and RAPD variation in 20 Swiss populations along five altitudinal gradients at four different elevations in the northern Swiss Alps. Additionally, allozyme and phenotypic variations in one Italian and two Spanish populations were compared with the variation in the Swiss populations. We hypothesized that there will be statistically significant genetic differences among populations of different altitudes and sites. The results showed no substantial correlation between genetic variation and phenotypic variation among Swiss populations. These results imply that outbreeding and effective gene exchange (long-distance spore dispersal) are the keys to population structure in this fern species, and as a consequence, phenotypic plasticity is assumed to be favored. This contrasts with results found in similar studies of herbaceous flowering plants where genetic adaptation to gradients like altitude is common. However, when data from the more distant Italian and Spanish populations of A. filix-femina were included, significant variation was detected.     

Adaptive divergence in plasticity in natural populations of Impatiens capensis and its consequences for performance in novel habitats

We tested for adaptive differentiation between two natural populations of Impatiens capensis from sites known to differ in selection on plasticity to density. We also determined the degree to which plasticity to density within a site was correlated with plastic responses of experimental immigrants to foreign sites. Inbred lines, derived from natural populations in an open-canopy site and a woodland site, were planted reciprocally in both original sites at naturally occurring high densities and at low density. The density manipulation represents environmental variation typically experienced within the site of a given population, and the transplant manipulation represents environmental differences between sites of different populations. Internode elongation, meristem allocation, leaf length, flowering date, and total lifetime fitness were measured. Genotypes originating in the open site, where selection favored plasticity of first internode length and flowering time (Donohue et al. 2000a), were more plastic in those characters than genotypes originating from the woodland site, where plasticity was maladaptive. Therefore, these two populations appear to have responded to divergent selection on plasticity. Plasticity to density strongly resembled plasticity to site differences for many characters, suggesting that similar environmental factors elicit plasticity both to density and to overhead canopy. Thus, plasticity that evolved in response to density variation within a site influenced phenotypic expression in the foreign site. Plastic responses to site caused immigrants from foreign populations to resemble native genotypes more closely. In particular, immigrants from the open site converged toward the selectively favored early-flowering phenotype of native genotypes in the woodland site, thereby reducing potential fitness differences between foreign and native genotypes. However, because genotypes from the woods population were less plastic than genotypes from the sun population, phenotypic differences between populations were greatest in the open site at low density. Therefore, population differences in plasticity can cause genotypes from foreign populations to be more strongly selected against in some environments than in others. However, genetic constraints and limits to plasticity prevented complete convergence of immigrants to the native phenotype in any environment.     

Phenotypic plasticity inLemna minor (Lemnaceae)

Eight genotypes ofLemna minor, originating from four continents, were grown for 15 days in eight different environmental treatments. Fronds under each treatment were then transferred into each of the eight environmental conditions for 15 days. The rate of frond production (relative growth rate) and mean frond biomass were recorded for each pre- and post-transfer treatment and root length was measured for each pre-transfer treatment. For all the traits, the levels of response varied significantly between genotypes (G) and between environmental conditions (E). G × E interaction effect was significant for all traits under pre-transfer treatments and some post-transfer treatments. Both pattern and amount of plasticity were genotypically variable but the amount of variation depended on the trait. The trait representing the best estimate of fitness, growth rate, exhibited the least amount of plasticity and on average, showed the most conservative pattern of plasticity. In contrast, the trait least related to fitness, root length, was the most plastic and showed the most divergent pattern of plasticity. Under some post-transfer treatments, growth rate and mean frond biomass were affected by origin (initial treatment) effect. Pattern and amount of plasticity were also influenced by initial treatments. Since some genotypes may be more affected than others by environmental conditions, origin effect may accentuate G × E interaction and therefore, modify the pattern and amount of plasticity. Comparison between dendrograms based on genetic and phenotypic similarities suggested that there is no relationship between genetic and phenotypic divergence. This lack of relationship may be due to the fact that plasticity is not necessarily adaptive.     

Plasticity to simulated shade is associated with altitude in structured populations of Arabidopsis thaliana

Plants compete for photosynthesis light and induce a shade avoidance syndrome (SAS) that confers an important advantage in asymmetric competition for light at high canopy densities. Shade plasticity was studied in a greenhouse experiment cultivating Arabidopsis thaliana plants from 15 populations spread across an altitudinal gradient in the northeast area of Spain that contain a high genetic variation into a reduced geographical range. Plants were exposed to sunlight or simulated shade to identify the range of shade plasticity. Fourteen vegetative, flowering and reproductive traits were measured throughout the life cycle. Shade plasticity in flowering time and dry mass was significantly associated with the altitude of population origin. Plants from coastal populations showed higher shade plasticity indexes than those from mountains. The altitudinal variation in flowering leaf plasticity adjusted negatively with average and minimum temperatures, whereas dry mass plasticity was better explained by negative regressions with the average, maximum and minimum temperatures, and by a positive regression with average precipitation of the population origin. The lack of an altitudinal gradient for the widest number of traits suggests that shade light could be a driver explaining the distribution pattern of individuals in smaller geographical scales than those explored here.     

No effect of elevation and fragmentation on genetic diversity and structure in Polylepis australis trees from central Argentina

Phenological differences in flowering arising along elevational gradients may be caused by either local adaptation or phenotypic plasticity. Local adaptation can lead to reproductive isolation of populations at different elevational zones and thus produce elevational genetic structuring, while phenotypic plasticity does not produce elevational genetic structuring. In this study, we examined the effects of elevation and fragmentation on genetic diversity and structure of Polylepis australis populations, where individuals exhibit phenological differences in flowering along an elevational gradient. We assessed the polymorphism of amplified fragment length polymorphism markers in adults and saplings from one conserved and one fragmented forest covering elevations from 1600 to 2600 m asl. Over 98% of variation was found within populations, and we found very low and similar genetic differentiation along elevational gradients for adults and saplings in both continuous and fragmented forests. In addition, there was no significant relationship between genetic diversity and elevation. Results indicated that phenological differences along elevational gradients are more likely caused by phenotypic plasticity than local adaptation, and fragmentation does not appear to have affected genetic diversity and differentiation in the studied populations. Results therefore imply that if necessary, seeds for reforestation purposes may be collected from different elevations to the seeding or planting sites.     

POPULATION AND GENOTYPE NICHE WIDTH IN CLONAL PHLOX PANICULATA

We use a model population comprised of five genotypes of Phlox paniculata L. to investigate the contribution of individuals to the response breadth (niche) of the population on a light gradient and a moisture gradient. Analysis of within- and between-genotype components of population response width showed up to 20% of the response is due to between-genotype effects, depending upon the character considered. Since the only way a sedentary organism can deal with a variable environment is through plasticity of response, differences in levels of phenotypic plasticity between genotypes on the two resource gradients were also investigated. There was no correlation between level of phenotypic plasticity and success over a range of environments. Niche breadth calculated as Levins' (B) and Roughgarden's (w²) indicated that flowering, and hence sexual reproduction, was limited to a much narrower range of environments than was vegetative growth. We also found significant genotype × environment interactions on both gradients, indicating differences in genetic response to the environment.     

Ecotype differentiation in Capsella

Seed samples were collected from wild populations of Capsella bursa-pastoris growing along a cline from low elevations to the high mountain region in Switzerland and from different latitudes in Scandinavia.Progeny were grown in open-field random block experiments, in transplantation experiments and in growth chambers. Beginning of flowering was recorded. Under various environmental conditions we got rough ecotypic complexes: Early flowering Scandinavian populations and late flowering populations from the Alps. A quantitative promotion in a larger photoperiod exists in all populations (quantitative LTP).Along various considered altitude gradients in the Alps populations from higher elevations are later flowering and tend to overwinter (biennial).Obviously there exists a very distinct adaptation in graduate ecotypical differentiation. But geographical and climatical adaptation could be superimposed by local human influences like agriculture. The physiological and genetical background is present, but until now nearly invisible.     

Experimental alteration of DNA methylation affects the phenotypic plasticity of ecologically relevant traits in <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

Heritable phenotypic variation in plants can be caused not only by underlying genetic differences, but also by variation in epigenetic modifications such as DNA methylation. However, we still know very little about how relevant such epigenetic variation is to the ecology and evolution of natural populations. We conducted a greenhouse experiment in which we treated a set of natural genotypes of Arabidopsis thaliana with the demethylating agent 5-azacytidine and examined the consequences of this treatment for plant traits and their phenotypic plasticity. Experimental demethylation strongly reduced the growth and fitness of plants and delayed their flowering, but the degree of this response varied significantly among genotypes. Differences in genotypes’ responses to demethylation were only weakly related to their genetic relatedness, which is consistent with the idea that natural epigenetic variation is independent of genetic variation. Demethylation also altered patterns of phenotypic plasticity, as well as the amount of phenotypic variation observed among plant individuals and genotype means. We have demonstrated that epigenetic variation can have a dramatic impact on ecologically important plant traits and their variability, as well as on the fitness of plants and their ecological interactions. Epigenetic variation may thus be an overlooked factor in the evolutionary ecology of plant populations.     

Natural selection contributes to geographic patterns of thermal plasticity in Plantago lanceolata

A long‐standing debate in evolutionary biology concerns the relative importance of different evolutionary forces in explaining phenotypic diversification at large geographic scales. For example, natural selection is typically assumed to underlie divergence along environmental gradients. However, neutral evolutionary processes can produce similar patterns. We collected molecular genetic data from 14 European populations of Plantago lanceolata to test the contributions of natural selection versus neutral evolution to population divergence in temperature‐sensitive phenotypic plasticity of floral reflectance. In P. lanceolata, reflectance plasticity is positively correlated with latitude/altitude. We used population pairwise comparisons between neutral genetic differentiation (F_ST and Jost's D) and phenotypic differentiation (P_ST) to assess the contributions of geographic distance and environmental parameters of the reproductive season in driving population divergence. Data are consistent with selection having shaped large‐scale geographic patterns in thermal plasticity. The aggregate pattern of P_ST versus F_ST was consistent with divergent selection. F_ST explained thermal plasticity differences only when geographic distance was not included in the model. Differences in the extent of cool reproductive season temperatures, and not overall temperature variation, explained plasticity differences independent of distance. Results are consistent with the hypothesis that thermal plasticity is adaptive where growing seasons are shorter and cooler, that is, at high latitude/altitude.     

Effects of endophytic fungi on the phenotypic plasticity of Lolium perenne (Poaceae)

The effects of high vs. low levels of endophytic fungi on the phenotypic plasticity of cloned genotypes were examined in perennial ryegrass (Lolium perenne L.). The objectives were to determine whether endophytic fungi influence plastic responses of host genotypes to variable soil nutrients and whether or not endophyte infection and host genotype interact to determine the extent of this plasticity. Twelve infected genotypes were cloned into ramets: half the ramets were treated with the systemic fungicide Benomyl to reduce or eliminate the endophyte, while the other half were untreated. Ramets of each genotype were subjected to high, medium, or low levels of nutrients in the greenhouse for 11 wk. Tiller number, leaf area, and leaf mass were determined after 11 and 25 wk. The fungicide significantly reduced the level of endophyte infection. Responses to nutrient conditions in relation to fungicide treatment were genotype specific: for some genotypes, high levels of endophytic fungi appeared to reduce plasticity, while for other genotypes the endophyte had no effect. The potential for microscopic symbionts to affect phenotypic plasticity in genetically variable populations has not often been recognized. However, the clandestine effects of symbionts on the plasticity of host genotypes could impact microevolutionary processes occurring within plant populations that occupy heterogeneous environments.     

Phenotypic plasticity in Cardamine flexuosa: variation among populations in plastic response to chilling treatments and photoperiods

Variability in the plastic responses of seven life history traits to different chilling and photoperiod regimes was studied in four wild populations of Cardamine flexuosa. This species, a winter-green or year-long annual, showed a facultative long-day and chilling requirement for flowering. Considerable variation among populations was noted in plasticity of all traits. Differences in plasticity were greater among three paddy field populations from different climatic areas than between adjacent populations under different disturbance regimes. A paddy field population (OP) and an adjacent orchard population (OG) exhibited similar plasticity, in both amounts and patterns of response. TP, a Japan Sea coast population, was distinct from three other populations, especially in the small amounts of plasticity. Differences in amount of response were much more common than differences in pattern of response. Character expressions of five traits were significantly correlated with the number of days to flowering. Days to flowering and the numbers of inflorescences and siliques showed high negative correlations because the branching ability of meristems decreased with delay of flowering.     

Correlation between time of flowering and phenotypic plasticity in Arabidopsis thaliana (Brassicaceae)

We observed substantial variation in the time of flowering among 13 populations of Arabidopsis thaliana (Brassicaceae) from an extensive latitudinal range when grown under uniform experimental conditions. The later the onset of flowering, the greater was potential reproduction. Later flowering plants also had greater plasticity in a host of morphological and physiological traits measured in nutrient-rich vs. nutrient-poor test environments. This relationship between flowering time and overall plasticity was only apparent for traits measured at the time of seed production, not at the time of flowering or earlier. At the time of seed production in this short-lived annual, the regression of a multivariate measure of overall plasticity on the time of flowering was linear and highly significant (r² = 0.90, P < 0.0001). These correlations among time of flowering, reproductive fitness, and plasticity support the idea that selection for late-flowering genotypes would select concomitantly for greater plasticity.     

Bergmann's rule is maintained during a rapid range expansion in a damselfly

Climate‐induced range shifts result in the movement of a sample of genotypes from source populations to new regions. The phenotypic consequences of those shifts depend upon the sample characteristics of the dispersive genotypes, which may act to either constrain or promote phenotypic divergence, and the degree to which plasticity influences the genotype–environment interaction. We sampled populations of the damselfly Erythromma viridulum from northern Europe to quantify the phenotypic (latitude–body size relationship based on seven morphological traits) and genetic (variation at microsatellite loci) patterns that occur during a range expansion itself. We find a weak spatial genetic structure that is indicative of high gene flow during a rapid range expansion. Despite the potentially homogenizing effect of high gene flow, however, there is extensive phenotypic variation among samples along the invasion route that manifests as a strong, positive correlation between latitude and body size consistent with Bergmann's rule. This positive correlation cannot be explained by variation in the length of larval development (voltinism). While the adaptive significance of latitudinal variation in body size remains obscure, geographical patterns in body size in odonates are apparently underpinned by phenotypic plasticity and this permits a response to one or more environmental correlates of latitude during a range expansion.     

Two reproductive traits show contrasting genetic architectures in Plantago lanceolata

In many species, temperature‐sensitive phenotypic plasticity (i.e., an individual's phenotypic response to temperature) displays a positive correlation with latitude, a pattern presumed to reflect local adaptation. This geographical pattern raises two general questions: (a) Do a few large‐effect genes contribute to latitudinal variation in a trait? (b) Is the thermal plasticity of different traits regulated pleiotropically? To address the questions, we crossed individuals of Plantago lanceolata derived from northern and southern European populations. Individuals naturally exhibited high and low thermal plasticity in floral reflectance and flowering time. We grew parents and offspring in controlled cool‐ and warm‐temperature environments, mimicking what plants would encounter in nature. We obtained genetic markers via genotype‐by‐sequencing, produced the first recombination map for this ecologically important nonmodel species, and performed quantitative trait locus (QTL) mapping of thermal plasticity and single‐environment values for both traits. We identified a large‐effect QTL that largely explained the reflectance plasticity differences between northern and southern populations. We identified multiple smaller‐effect QTLs affecting aspects of flowering time, one of which affected flowering time plasticity. The results indicate that the genetic architecture of thermal plasticity in flowering is more complex than for reflectance. One flowering time QTL showed strong cytonuclear interactions under cool temperatures. Reflectance and flowering plasticity QTLs did not colocalize, suggesting little pleiotropic genetic control and freedom for independent trait evolution. Such genetic information about the architecture of plasticity is environmentally important because it informs us about the potential for plasticity to offset negative effects of climate change.     

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.     

Developmental thermal plasticity among Drosophila melanogaster populations

Many biotic and abiotic variables influence the dispersal and distribution of organisms. Temperature has a major role in determining these patterns because it changes daily, seasonally and spatially, and these fluctuations have a significant impact on an organism's behaviour and fitness. Most ecologically relevant phenotypes that are adaptive are also complex and thus they are influenced by many underlying loci that interact with the environment. In this study, we quantified the degree of thermal phenotypic plasticity within and among populations by measuring chill‐coma recovery times of lines reared from egg to adult at two different environmental temperatures. We used sixty genotypes from six natural populations of Drosophila melanogaster sampled along a latitudinal gradient in South America. We found significant variation in thermal plasticity both within and among populations. All populations exhibit a cold acclimation response, with flies reared at lower temperatures having increased resistance to cold. We tested a series of environmental parameters against the variation in population mean thermal plasticity and discovered the mean thermal plasticity was significantly correlated with altitude of origin of the population. Pairing our data with previous experiments on viability fitness assays in the same populations in fixed and variable environments suggests an adaptive role of this thermal plasticity in variable laboratory environments. Altogether, these data demonstrate abundant variation in adaptive thermal plasticity within and among populations.     

Phytotron experiments in Pisum

Summary The flowering behaviour of 17 Pisum mutants and 20 recombinants was studied under three different temperatures using long-day phytotron conditions. A constant low temperature of 12.5 ° C led to a strong delay in flowering in all the genotypes tested but distinct relative differences could be found between them. Relative differences were also present with regard to speed of ontogenetic development under a permanent high temperature of 25.5 °C or under an alternating change between low and high temperature. Under the low temperature, recombinants R 20D and R 20E, carrying gene efr for earliness, entered the flowering period more than 4 weeks later than the donor of efr, demonstrating thereby a negative influence of one of the other mutant genes on efr. The high temperature of 25 °C influenced the flowering behaviour of 4 fasciated genotypes negatively — in contrast to the other strains studied. The plants of recombinant R 405 produced only tiny flower buds under these conditions. None of the plants of recombinant R 142F flowered under either the constant low or high temperature — they need the change of low and higher temperature for normal flower formation. The experiments show that most of the genotypes tested react specifically to the three temperature conditions offered to them.