Potential for selection on plants for water-use efficiency as estimated by carbon isotope discrimination

Water-use efficiency is thought to be related to plant performance and natural selection for plants in arid habitats, based on a general expectation that increased water-use efficiency is associated with decreased carbon gain and biomass accumulation. Using leaf carbon isotope discrimination Δ to determine integrated water-use efficiency, we estimated genetic variance for, and examined the relationships among Δ, biomass, and gas exchange characters for full-sibling families of the woody shrub, Chrysothamnus nauseosus, grown from seed collected at Tintic, Utah. In both well-watered greenhouse and common garden experiments, and water-limited common garden experiments, there were significant family differences for Δ, biomass, and morphological characters, indicating a potential for genetic change in response to selection. However, estimates of broad-sense heritabilities for Δ were low, indicating that the rate of change in response to selection would be relatively slow. This was consistent with the large amount of phenotypic plasticity observed for Δ as it differed with water treatment and year in the garden experiment. Phenotypically, aboveground biomass and Δ were negatively correlated within the well-watered treatments (i.e., more water-use efficient plants were larger), not correlated within the water-limited treatment, and positively correlated for combined well-watered and water-limited garden treatments, suggesting that variation in both photosynthetic capacity and stomatal limitation contribute to the variation in Δ. In contrast to the phenotypic correlations, genetic correlations for biomass and Δ were consistently negative within each treatment, and selection for higher water-use efficiency through low Δ for C. nauseosus plants in this population would tend to shift populations toward larger plants. For C. nauseosus, increased water-use efficiency is not necessarily associated with decreased carbon gain.     

Phenotypic plasticity in Phlox

Summary Three species of Phlox (Polemoniaceae) were grown in 6 greenhouse treatments. A variety of traits were recorded and the correlations among them were computed for each treatment. The phenotypic correlations between characters are significantly altered when plants are grown under different environmental conditions. These changes in correlation structure result from the differential phenotypic plasticity of traits. Partial correlations between flower production and other traits are also environment-dependent. Such changes can alter the intensity of, and possibly the response to, selection on traits correlated with fitness in natural plant populations.     

Phenotypic plasticity in host choice behavior in black bean aphid, Aphis fabae (Homoptera: Aphididae)

The study of phenotypic plasticity in host choice behavior is crucial to predict evolutionary patterns of insect–plant interactions. The presence of sufficient variation in plasticity may facilitate host race formation and sympatric speciation. In this study, 13 Aphis fabae Scopoli genotypes reared both on broad bean and nasturtium exhibited statistically significant genotypic variability in host selection behavior. Some genotypes displayed increase in preference and acceptance in a novel host plant through generations. There are also strong conditioning effects of nasturtium as nasturtium reared genotypes are more willing to choose nasturtium over broad bean while broad bean reared genotypes do not show differences in choosing between the two host plants. There are also positive relationships between fitness and host choice behavior particularly for nasturtium. Results of the study supported the hypothesis that phenotypic plasticity in host choice behavior may be one of the major determinants of the evolutionary trajectory of a parasitic species, such as aphids.     

Phenotypic plasticity for skeletal growth,density and calcification of <Emphasis Type="Italic">Porites lobata</Emphasis> in response to habitat type

A reciprocal transplant experiment (RTE) of the reef-building coral Porites lobata between shallow (1.5 m at low tide) back reef and forereef habitats on Ofu and Olosega Islands, American Samoa, resulted in phenotypic plasticity for skeletal characteristics. Transplants from each source population (back reef and forereef) had higher skeletal growth rates, lower bulk densities, and higher calcification rates on the back reef than on the forereef. Mean annual skeletal extension rates, mean bulk densities, and mean annual calcification rates of RTE groups were 2.6–9.8 mm year⁻¹, 1.41–1.44 g cm⁻³, and 0.37–1.39 g cm⁻² year⁻¹ on the back reef, and 1.2–4.2 mm year⁻¹, 1.49–1.53 g cm⁻³, and 0.19–0.63 g cm⁻² year⁻¹ on the forereef, respectively. Bulk densities were especially responsive to habitat type, with densities of transplants increasing on the high energy forereef, and decreasing on the low energy back reef. Skeletal growth and calcification rates were also influenced by source population, even though zooxanthella genotype of source colonies did not vary between sites, and there was a transplant site x source population interaction for upward linear extension. Genetic differentiation may explain the source population effects, or the experiment may have been too brief for phenotypic plasticity of all skeletal characteristics to be fully expressed. Phenotypic plasticity for skeletal characteristics likely enables P. lobata colonies to assume the most suitable shape and density for a wide range of coral reef habitats.     

DIFFERING SELECTION ON PLANT PHYSIOLOGICAL TRAITS IN RESPONSE TO ENVIRONMENTAL WATER AVAILABILITY: A TEST OF ADAPTIVE HYPOTHESES

I used phenotypic selection analysis to test the prediction from functional and comparative studies of plants that smaller leaves and more efficient water use are adaptive in drier environments. I measured selection gradients on leaf size and instantaneous water-use efficiency (a measure of carbon gain per unit water loss) in experimental populations of Cakile edentula var. lacustris placed into wet and dry environments in the field. Linear and nonlinear selection differed significantly between the two environments as predicted. Water-use efficiency was selected to be higher, and leaf area was selected toward a small intermediate optimum, in the dry environment. There was also significant positive correlational selection on water-use efficiency and leaf size, suggesting that the optimum leaf size in the dry environment is greater for plants with higher water-use efficiency. In contrast, neither leaf size nor water-use efficiency were selected in the wet environment, though larger leaves resulted in greater vegetative biomass. Path analysis of the linear selection gradients found that water-use efficiency affected plant fitness primarily because it increased vegetative biomass, as suggested by the hypotheses about the function of physiological traits. These results were not only consistent with the functional hypotheses but also with the observed genetic differentiation in water-use efficiency and leaf size between wet and dry site populations.     

Plastic bet-hedging in an amphicarpic annual: an integrated strategy under variable conditions

Amphicarpy is a form of diversified bet-hedging expressed mostly in annual plants, where two types of offspring are produced with two distinct ecological roles: long-range aerial dispersers and highly competitive subterranean, sedentary fruit. Emex spinosa is a semi-arid, amphicarpic annual, inhabiting habitats with different levels of environmental variation. We tested the hypothesis that, in E. spinosa, bet-hedging may be “fine-tuned” by plasticity in the phenotype ratio (aerial/subterranean fruit mass) as a function of environmental conditions. We conducted a greenhouse experiment, manipulating nutrient availability and intraspecific density, to determine the pattern of ratio shifts. In order to determine whether the integrated strategy is an adaptation to variable habitats, a similar common garden experiment was conducted, comparing two natural populations differing in environmental variability. The offspring ratio shifted in response to both nutrient availability and plant density. In pots containing single plants the ratio increased steeply with nutrient availability, while in pots containing eight plants a more moderate increase occurred. These shifts were the result of plasticity in allocation to both achene types, as well as ontogenetic effects on aerial achene production. The degree of response increased with the heterogeneity of the habitat of origin. We found evidence for an adaptive integrated strategy, with bet-hedging “fine-tuned” by phenotypic plasticity. Strenuous conditions tended to shift the offspring ratio towards securing subterranean reproductive success, while favorable conditions resulted in a shift towards dispersible achenes. The authors Asaf Sadeh and Hagai Guterman contributed equally to this study.     

Consumption rates and the evolution of diet-induced plasticity in the head morphology of Melanoplus femurrubrum (Orthoptera: Acrididae)

Summary Phenotypic plasticity may be an ecologically important evolutionary response to natural selection in multiple environments. I have determined the effect of diet-induced developmental plasticity in the head size of grasshoppers (Melanoplus femurrubrum) onfeeding performance on two types of plants. Full-sib families were divided and raised on either red clover, Trifolium repens, or rye grass, Lolium perenne. In three different stages of ontogeny, grasshoppers raised on rye grass had significantly larger heads, relative to body size, than full-sibs raised on clover. A principal components analysis indicated that two to five relative head size characters covaried as a block in their plastic response to the feeding environment. Regressions of adjusted consumption rates (mg/sec) against relative head size revealed that larger head sizes, induced by the rye grass diet, enhanced consumption rates of rye grass, but not clover. Unexpectedly, a similar positive association was observed between head size and consumption rate for grasshoppers raised on clover when they were feeding on clover. These results support the inference that grasshoppers exhibit adaptive phenotypic plasticity. However, the unexpected influence of head size on consumption rates of clover indicates that the functional relationship between head morphology and feeding performance is complex and that variation in this relationship among plant environments is not sufficient to explain the evolution of diet-induced phenotypic plasticity.     

Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna

In frequently burnt mesic savannas, trees can get trapped into a cycle of surviving fire-induced stem death (i.e. topkill) by resprouting, only to be topkilled again a year or two later. The ability of savanna saplings to resprout repeatedly after fire is a key component of recent models of tree–grass coexistence in savannas. This study investigated the carbon allocation and biomass partitioning patterns that enable a dominant savanna tree, Acacia karroo, to survive frequent and repeated topkill. Root starch depletion and replenishment, foliage recovery and photosynthesis of burnt and unburnt plants were compared over the first year after a burn. The concentration of starch in the roots of the burnt plants (0.08 ± 0.01 g g⁻¹) was half that of the unburnt plant (0.16 ± 0.01 g g⁻¹) at the end of the first growing season after topkill. However, root starch reserves of the burnt plants were replenished over the dry season and matched that of unburnt plants within 1 year after topkill. The leaf area of resprouting plants recovered to match that of unburnt plants within 4–5 months after topkill. Shoot growth of resprouting plants was restricted to the first few months of the wet season, whereas photosynthetic rates remained high into the dry season, allowing replenishment of root starch reserves. ¹⁴C labeling showed that reserves were initially utilized for shoot growth after topkill. The rapid foliage recovery and the replenishment of reserves within a single year after topkill implies that A. karroo is well adapted to survive recurrent topkill and is poised to take advantage of unusually long fire-free intervals to grow into adults. This paper provides some of the first empirical evidence to explain how savanna trees in frequently burnt savannas are able to withstand frequent burning as juveniles and survive to become adults. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variation in and adaptive plasticity of flower size and drought-coping traits

Although phenotypic plasticity of morphological and physiological traits in response to drought could be adaptive, there have been relatively few tests of plasticity variation or of adaptive plasticity in drought-coping traits across populations with different moisture availabilities. We measured floral size, vegetative size, and physiological traits in four field populations of Leptosiphon androsaceus (Polemoniaceae) that were distributed across a rainfall gradient in California, USA. Measurements were made over 5 years that varied in precipitation. We also conducted a growth chamber experiment in which half-sibs from three populations were divided equally among a well-watered and a drought treatment. We tested for selection on traits in each of the watering treatments, and evaluated whether traits exhibited plasticity. In the field, plant traits exhibited substantial variation across populations and years. Flower size, leaf size, and water-use efficiency (WUE) were generally higher for populations that received greater average rainfall. However, in dry years, we observed a decrease in flower and leaf size, but an increase in WUE across the populations. In the growth chamber experiment, leaf and physiological traits exhibited plasticity, with smaller leaves and higher WUE found in the drought, as compared to the well-watered treatment. Only specific leaf area exhibited differentiation in plasticity among populations. Although there was no observed plasticity in floral size, selection favored smaller flowers in the drought treatment and larger flowers in the well-watered treatment. Our results suggest that moisture availability has led to trait variation in L. androsaceus via a combination of selection and phenotypic plasticity.     

<Emphasis Type="Italic">In vitro</Emphasis> selection and regeneration of chrysanthemum (<Emphasis Type="Italic">Dendranthema grandiflorum</Emphasis> Tzelev) plants resistant to culture filtrate of <Emphasis Type="Italic">Septoria obesa</Emphasis> Syd

Callus cultures derived from leaf segments of chrysanthemum cultivar ‘Snow Ball’ which was susceptible to Septoria obesa were successfully used for in vitro selection for resistance to this pathogenic fungus. Resistant cell lines were selected by culturing callus on growth medium containing various concentrations of S. obesa filtrate. Resistant calluses obtained after two cycles (30 d each cycle) of selection were used for plant regeneration. About 30% of the plants regenerated from the resistant calluses and 70–80% of the plants raised from cuttings had acquired considerable resistance against the pathogen in the field. No phenotypic variation was observed in the selected regenerates.     

Comparisons of carbon isotope discrimination in populations of aridland plant species differing in lifespan

Summary Carbon isotope discrimination () was compared between populations of dominant perennial plant species, differing in life expectancy, in two deserts with contrasting vegetation types. In both deserts, plants of the shorter-lived species showed significantly higher and greater intrapopulation variance in this character compared to the long-lived species. These results indicate underlying differences in gas-exchange physiology, and suggest a positive correlation between water-use efficiency and lifespan in desert plants. Differences in variance for this character may reflect greater microenvironmental variation experienced by shorter-lived plants and/or different forms of selection acting on water-use traits. Spatial distributions were significantly clustered for the shorter-lived species and significantly uniform for the long-lived species, indicating that competition has been important in the development of the long-lived populations. The long-lived Larrea tridentata showed a significant, negative correlation between and Thiessen polygon area, suggesting a positive relationship between water-use efficiency and longevity within this species. This relationship was weakly supported in the other warm desert species, Encelia farinosa, but was not observed within populations of the cold desert species, Gutierrezia microcephala and Coleogyne ramosissima. These results suggest that reflects key aspects of plant metabolism related to lifespan; these differences may ultimately influence interactions among desert plants and the structure of desert plant communities.     

<Emphasis Type="Italic">Lotus corniculatus</Emphasis> L. response to carbon dioxide concentration and radiation level variations

Carbon dioxide concentration and light conditions may greatly vary between mountainous and lowland areas determining the photosynthetic performance of plants species. This paper aimed to evaluate the photosynthetic responses of Lotus corniculatus, growing in a mountain and a lowland grassland, under low and high radiation and CO₂ concentration. Net photosynthetic rate, stomatal conductance, transpiration rate, and intercellular CO₂ concentration were measured while the water-use efficiency and the ratio of variable to maximal fluorescence were calculated. Photosynthetic response curves to different levels of radiation and intercellular CO₂ partial pressure were estimated. Our results showed that high radiation and CO₂ concentration enhanced water-use efficiency of plants at both sites, enabling them to use more efficiently the available water reserves under drought conditions. The increase of radiation and CO₂ concentration would enhance the photosynthetic performance of the mountainous population of L. corniculatus, which overall seems to express higher phenotypic plasticity.     

Phenotypic plasticity of Capsella bursa-pastoris (Brassicaceae) and its effect on fitness in response to temperature and soil moisture

Capsella bursa-pastoris is one of the most common plants on earth. Although phenotypic plasticity of ecologically important traits possibly contributes to its wide geographic range, little is known about the plasticity of C. bursa-pastoris and its effects on its fitness. In a laboratory, we assessed the phenotypic plasticity in response to two representative climatic conditions: temperature and soil moisture. In addition, we quantitatively evaluated the relationship between phenotypic plasticity and fruit production. Most measured morphological and physiological traits exhibited plastic responses to temperature and fitness based on fruit production was maintained across temperatures. In contrast, no plasticity to soil moisture was detected, and plants produced fewer fruits in dry soil. Selection analysis revealed that the plasticity of the flowering time and the water-use efficiency had positive effects on fruit production over the tested temperature regime. These experimental results suggest that phenotypic plasticity probably enables C. bursa-pastoris to cope with heterogeneous temperature environments and thereby probably contributes to its wide geographic range.     

Carbon isotope composition of N2-fixing and N-fertilized legumes along an elevational gradient

Dinitrogen-fixing legumes are frequently assumed to be less water-use efficient than plants utilizing soil mineral N, because of the high respiratory requirements for driving N₂ fixation. However, since respiration is assumed not to discriminate against ¹³C, any differences in water-use efficiency exclusively due to respiration should not be apparent in carbon isotope discrimination () values. Our objective was to determine if the source of N (N₂ fixation versus soil N) had any effect on of field-grown grain legumes grown at different elevations. Four legume species, Glycine max, Phaseolus lunatus, P. vulgaris, and Vigna unguiculata, were grown on five field sites spanning a 633 m elevational gradient on the island of Maui, Hawaii. The legumes were either inoculated with a mixture of three effective strains of rhizobia or fertilized weekly with urea at 100 kg N ha^-1 in an attempt to completely suppress symbiotic N₂-fixing activity. In 14 of 20 analyses of stover and 12 of 15 analyses of seed values were significantly higher (p=0.10) in the inoculated plants than the N-fertilized plants. Nitrogen concentrations were generally higher in the fertilized treatments than the inoculated treatments. The different values obtained depending on N-source may have implications in using as an indicator of water-use efficiency or yield potential of legumes.     

Phenotypic plasticity of invasive Alternanthera philoxeroides in relation to different water availability, compared to its native congener

Phenotypic plasticity and genetic differentiation are two possible mechanisms that plants use to cope with varying environments. Although alligator weed (Alternanthera philoxeroides) possesses very low genetic diversity, this alien weed has successfully invaded diverse habitats with considerably varying water availability (from swamps to dry lands) in China. In contrast, its native congener (Alternanthera sessilis) has a much narrower ecological breadth, and is usually found in moist habitats. To understand the mechanisms underlying the contrasting pattern, we performed a greenhouse experiment to compare the reaction norms of alligator weed with those of its native congener, in which water availability was manipulated. Our results revealed that the two congeners had similar direction of phenotypic plasticity. However, A. philoxeroides showed greater plasticity in amount than did A. sessilis in many traits examined during the switch from wet to drought treatment. Nearly all of the phenotypic variance in A. philoxeroides could be ascribed to plasticity, while A. sessilis had a much higher fraction of phenotypic variance that could be explained by genotypic variation. These interspecific differences in plastic responses to variable water availability partially explained the difference in spatial distribution of the two congeners.     

Cloning of a Novel Omega-6 Desaturase from Flax (<Emphasis Type="Italic">Linum usitatissimum</Emphasis> L.) and Its Functional Analysis in <Emphasis Type="Italic">Saccharomyces cerevisiae</Emphasis>

The Δ12 desaturase represents a diverse gene family in plants and is responsible for conversion of oleic acid (18:1) to linoleic acid (18:2). Several members of this family are known from plants like Arabidopsis and Soybean. Using primers from conserved C- and N-terminal regions, we have cloned a novel Δ12 desaturase gene amplified from flax genomic DNA, denoted as LuFAD2-2. This intron-less gene is 1,149-base pair long encoding 382 amino acids—putative membrane-bound Δ12 desaturase protein. Sequence comparisons show that the novel sequence has 85% similarity with previously reported flax Δ12 desaturase at amino acid level and shows typical features of membrane-bound desaturase such as three conserved histidine boxes along with four membrane-spanning regions that are universally present among plant desaturases. The signature amino acid sequence ‘YNNKL’ was also found to be present at the N terminus of the protein, which is necessary and sufficient for ER localization of enzyme. Neighbor-Joining tree generated from the sequence alignment grouped LuFAD2-2 among the other FAD2 sequences from Ricinus, Hevea, Jatropha, and Vernicia. When LuFAD2-2 and LuFAD2 were expressed in Saccharomyces cerevisiae, they could convert the oleic acid to linoleic acid, with an average conversion rate of 5.25 and 8.85%, respectively. However, exogenously supplied linoleic acid was feebly converted to linolenic acid suggesting that LuFAD2-2 encodes a functional FAD2 enzyme and has substrate specificity similar to LuFAD2.     

Phenotypic and transgenerational plasticity promote local adaptation to sun and shade environments

Adaptive plasticity is expected to be important when the grain of environmental variation is encompassed in offspring dispersal distance. We investigated patterns of local adaptation, selection and plasticity in an association of plant morphology with fine-scale habitat shifts from oak canopy understory to adjacent grassland habitat in Claytonia perfoliata. Populations from beneath the canopy of oak trees were >90 % broad leaved and large seeded, while plants from adjacent grassland habitat were >90 % linear-leaved and small seeded. In a 2-year study, we used reciprocal transplants and phenotypic selection analysis to investigate local adaptation, selection, plasticity and maternal effects in this trait-environment association. Transgenerational effects were studied by planting offspring of inbred maternal families grown in both environments across the same environments in the second year. Reciprocal transplants revealed local adaptation to habitat type: broad-leaved forms had higher fitness in oak understory and linear-leaved plants had higher fitness in open grassland habitat. Phenotypic selection analyses indicated selection for narrower leaves and lower SLA in open habitat, and selection for broad leaves and intermediate values of SLA in understory. Both plant morphs exhibited plastic responses in traits in the same direction as selection on traits (narrower leaves and lower SLA in open habitat) suggesting that plasticity is adaptive. We detected an adaptive transgenerational effect in which maternal environment influenced offspring fitness; offspring of grassland-reared plants had higher fitness than understory-reared plants when grown in grassland. We did not detect costs of plasticity, but did find a positive association between leaf shape plasticity and fitness in linear-leaved plants in grassland habitat. Together, these findings indicate that fixed differences in trait values corresponding to selection across habitat contribute to local adaptation, but that plasticity and maternal environmental effects may be favored through promotion of survival across heterogeneous environments.     

Trait Values,Not Trait Plasticity,Best Explain Invasive Species' Performance in a Changing Environment

The question of why some introduced species become invasive and others do not is the central puzzle of invasion biology. Two of the principal explanations for this phenomenon concern functional traits: invasive species may have higher values of competitively advantageous traits than non-invasive species, or they may have greater phenotypic plasticity in traits that permits them to survive the colonization period and spread to a broad range of environments. Although there is a large body of evidence for superiority in particular traits among invasive plants, when compared to phylogenetically related non-invasive plants, it is less clear if invasive plants are more phenotypically plastic, and whether this plasticity confers a fitness advantage. In this study, I used a model group of 10 closely related Pinus species whose invader or non-invader status has been reliably characterized to test the relative contribution of high trait values and high trait plasticity to relative growth rate, a performance measure standing in as a proxy for fitness. When grown at higher nitrogen supply, invaders had a plastic RGR response, increasing their RGR to a much greater extent than non-invaders. However, invasive species did not exhibit significantly more phenotypic plasticity than non-invasive species for any of 17 functional traits, and trait plasticity indices were generally weakly correlated with RGR. Conversely, invasive species had higher values than non-invaders for 13 of the 17 traits, including higher leaf area ratio, photosynthetic capacity, photosynthetic nutrient-use efficiency, and nutrient uptake rates, and these traits were also strongly correlated with performance. I conclude that, in responding to higher N supply, superior trait values coupled with a moderate degree of trait variation explain invasive species'' superior performance better than plasticity per se.     

Exotic plant communities shift water-use timing in a shrub-steppe ecosystem

Semiarid areas in the US have realized extensive and persistent exotic plant invasions. Exotics may succeed in arid regions by extracting soil water at different times or from different depths than native plants, but little data is available to test this hypothesis. Using estimates of root mass, gravimetric soil water, soil-water potential, and stable isotope ratios in soil and plant tissues, we determined water-use patterns of exotic and native plant species in exotic- and native-dominated communities in Washington State, USA. Exotic and native communities both extracted 12 ± 2 cm of water from the top 120 cm of soil during the growing season. Exotic communities, however, shifted the timing of water use by extracting surface (0–15 cm) soil water early in the growing season (i.e., April to May) before native plants were active, and by extracting deep (0–120 cm) soil water late in the growing season (i.e., June to July) after natives had undergone seasonal senescence. We found that δ ¹⁸O values of water in exotic annuals (e.g., −11.8 ± 0.4 ‰ for Bromus tectorum L.) were similar to δ ¹⁸O values of surface soil water (e.g., −13.3 ± 1.4 ‰ at −15 cm) suggesting that transpiration by these species explained early season, surface water use in exotic communities. We also found that δ ¹⁸O values of water in taprooted exotics (e.g., −17.4 ± 0.3 ‰ for Centaurea diffusa Lam.) were similar to δ ¹⁸O values of deep soil water (e.g., −18.4 ± 0.1 ‰ at −120 cm) suggesting that transpiration by these species explained late season, deep water use. The combination of early-season, shallow water-use by exotic winter-actives and late-season, deep water-use by taprooted perennials potentially explains how exotic communities resist establishment of native species that largely extracted soil water only in the middle of the growing season (i.e., May to June). Early season irrigation or the planting of natives with established root systems may allow native plant restoration.     

Compensation for floral herbivory in <Emphasis Type="Italic">Solanum carolinense</Emphasis>: identifying mechanisms of tolerance

While a plant’s capacity to tolerate damage by herbivores can be studied as a single trait, it is important to recognize that tolerance is generally a result of the combined action of several different traits. Here, we report on a pair of experiments to identify mechanisms for tolerating floral herbivory in Solanum carolinense, an andromonoecious perennial herb that regularly suffers from high levels of florivory. We measured the effect of actual and simulated florivory on host-plant fitness and assessed which plant traits exhibited plasticity in response to florivory. In addition, for each of nine plant genets, we calculated tolerance indices and determined which traits were genetically correlated with tolerance. Traits that served to help S. carolinense tolerate florivory in terms of sexual reproduction included initiating more inflorescences, aborting fewer buds prior to anthesis and fewer ovaries after fertilization, and increasing the ratio of perfect:male flowers. In addition, the greater the levels of florivory, the more the plants allocated to root growth, which may promote tolerance through greater potential future reproduction. The plant population contained significant genetic variation for tolerance itself and for nearly all of the putative tolerance mechanisms, which suggests that S. carolinense has the potential to evolve greater tolerance through a variety of different routes in response to natural selection.