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PubMed2001

A molecular phylogeny of Costaceae (Zingiberales).

The phylogenetic relationships of Costaceae, a tropical monocotyledonous family sister to the gingers (Zingiberaceae), were investigated with a combination of two chloroplast loci (the trnL-F locus, including the trnL intron, the 3'trnL exon, and the trnL-F intergenic spacer, and the trnK locus, including the trnK intron and the matK coding region) and one nuclear locus (ITS1-5.8s-ITS2). The resulting parsimony analysis of selected taxa that demonstrate the range of floral morphological variation in the family shows that the Cadalvena-type [corrected] floral morphology is ancestral to the group and that both Tapeinochilos species and a Monocostus + Dimerocostus clade represent recent divergences. The genus Costus is broadly paraphyletic but Costus subgenus Eucostus K. Schum. represents a large monophyletic radiation that is poorly resolved. Within this clade, secondary analyses suggest that pollination syndrome, traditionally used for taxonomic and classification purposes within the genus Costus, is a relatively plastic trait of limited phylogenetic utility. This represents the first detailed investigation into intrageneric and interspecific evolutionary relationships within the family Costaceae and presents some novel evolutionary trends with respect to floral morphology and biogeography.

Chloroplasts↗
PubMed2006

Grazing-induced changes in cell wall silicification in a marine diatom.

In aquatic environments, diatoms (Bacillariophyceae) constitute a central group of microalgae which contribute to about 40% of the oceanic primary production. Diatoms have an absolute requirement for silicon to build-up their silicified cell wall in the form of two shells (the frustule). To date, changes in diatom cell wall silicification have been only studied in response to changes in the growth environment, with consistent increase in diatom silica content when specific growth rates decrease under nutrient or light limitations. Here, we report the first evidence for grazing-induced changes in cell wall silicification in a marine diatom. Cells grown in preconditioned media that had contained both diatoms and herbivores are significantly more silicified than diatoms grown in media that have contained diatoms alone or starved herbivores. These observations suggest that grazing-induced increase in cell wall silicification can be viewed as an adaptive reaction in habitats with variable grazing pressure, and demonstrate that silicification in diatoms is not only a constitutive mechanical protection for the cell, but also a phenotypically plastic trait modulated by grazing. In turn, our results corroborate the idea that plant-herbivore interactions, beyond grazing sensu stricto, contribute to drive ecosystem structure and biogeochemical cycles in the ocean.

Adaptation, Physiological↗
PubMed2005

Partial selfing, ecological disturbance and reproductive assurance in an invasive freshwater snail.

Although reproductive assurance (RA) might play a central role in the evolution of the selfing rate, this hypothesis has never been seriously investigated in an hermaphroditic animal. We studied the mating system of the freshwater snail Physa acuta in which the availability of mating partners might be highly variable, because this species is an efficient colonizer occupying unstable habitats. A total of 11 populations differing in ecological disturbance regime (water level, openness) and snail densities were monitored over 2 years. The outcrossing rate was estimated in ca 10 families per population using microsatellite markers and the progeny-array approach. Components of fecundity and survival were recorded for each progeny. Predominant outcrossing (t(m)=0.94) was detected, with a few individuals (4%) purely selfing. The outcrossing rate did not explain among-family variation in fitness components. None of the predictions formulated under the RA hypothesis were verified: (i) selfing was related neither to disturbed habitats, nor to temporal density fluctuations, (ii) it was positively related to population density, (iii) it co-occurred with multiple paternity, and (iv) it did not induce delayed reproduction. Explanations for these negative results are discussed in light of other arguments supporting the RA hypothesis in P. acuta, as well as alternative theories explaining the occurrence of partial selfing, as either a genetically fixed or plastic trait.

Animals↗
PubMed2006

Quantitative and evolutionary biology of alternative splicing: how changing the mix of alternative transcripts affects phenotypic plasticity and reaction norms.

Alternative splicing (AS) of pre-messenger RNA is a common phenomenon that creates different transcripts from a single gene, and these alternative transcripts affect phenotypes. The majority of AS research has examined tissue and developmental specificity of expression of particular AS transcripts, how this specificity affects cell function, and how aberrant AS is related to disease. Few studies have examined quantitative between-individual variation in AS within a cell or tissue type, or in relation to phenotypes, but the results are compelling: quantitative variation in AS affects plastic traits such as stress, anxiety, fear, egg production, muscle performance, energetics and plant growth. Genomic analyses of AS are also at a nascent stage, but have revealed a number of significant evolutionary patterns. Growing knowledge of upstream genes and kinases that regulate AS provides the as-yet little explored potential to examine how these genes and pathways respond to environmental and genotype variables. Research in this area can provide glimpses of a labyrinth of genetic architectures that have rarely been considered in evolutionary and organismal biology, or in quantitative genetics. The scarcity of contribution to knowledge about AS from these fields is illustrated by the fact that heritability of quantitative variation in AS has not yet been determined for any gene in any organism. New research tactics that incorporate quantitative analyses of AS will allow organismal and evolutionary biologists to attain a fuller mechanistic understanding of many of the traits they study, and may lead to more rapid discovery of functionally important polymorphisms.

Alternative Splicing↗
PubMed2001

Molecular genetic markers provide no evidence for reproductive isolation among retreat building phenotypes of the net-spinning caddisfly Macrostemum carolina.

Larvae of the stream-dwelling, filter-feeding caddisfly Macrostemum carolina construct silken catchnets within protective retreats. In the Savannah River, M. carolina individuals make three different retreats, each with a distinct water entrance hole: (i) at the end of a silken tube; (ii) with a approximately 180 degrees silken backstop; and (iii) flush with the top of the retreat. To resolve whether these different retreats represent alternative behavioural phenotypes within a single panmictic population or fixed phenotypes within three genetically distinct populations or species, we compared the allele frequencies at three polymorphic nuclear loci (allozyme electrophoresis for Gpi, Mpi and Pgm) and the mitochondrial DNA (mtDNA) haplotype frequencies among individuals displaying the three retreat morphs. We also calculated pairwise exact tests of population differentiation using the allozyme and mtDNA allele frequencies. No significant genetic differentiation was detected among caddisflies exhibiting the different retreat morphs. Therefore, these morphs apparently represent a single panmictic population in the Savannah River. Consequently, additional study is required to assess whether this retreat polymorphism is a phenotypically plastic trait under conditional control, or is mediated by alternative alleles at a Mendelian gene or genes (or a combination of the two).

Animals↗
PubMed2003

Differentiation between subpopulations of a polychromatic damselfly with respect to morph frequencies, but not neutral genetic markers.

The damselfly, Nehalennia irene (Hagen), has two distinct female colour morphs. Individuals of one morph have male-like colouration and pattern (androchromes), whereas gynochromes are different from males and androchromes in these respects. In several damselflies, such female-limited polychromatism is attributable to a single genetic locus. We developed six polymorphic genetic markers, which were codominant, to test for genetic differentiation in N. irene, collected from two sites located 8 km from one another in eastern Ontario, Canada. Based on three censuses spanning a 10 year period (1992-2001), morph ratios differed consistently and significantly between these two sites. However, subpopulations at these sites were not genetically differentiated with respect to the putatively neutral markers. Our results suggest that site differences in morph ratios of female N. irene cannot be explained by genetic drift, but are consistent with spatially variable selection operating on different morphs, perhaps mediated by male density. Alternatively, morph type may be a plastic trait and cues for induction may differ between sites.

Animals↗
PubMed2002

Nitrogen availability alters the expression of carnivory in the northern pitcher plant, Sarracenia purpurea.

Atmospheric transport and deposition of nutrients, especially nitrogen, is a global environmental problem with well-documented consequences for ecosystem dynamics. However, monitoring nitrogen deposition is relatively expensive, monitoring stations are widely spaced, and estimates and predicted impacts of nitrogen deposition are currently derived from spatial modeling and interpolation of limited data. Ombrotrophic ("rain-fed") bogs are nutrient-poor ecosystems that are especially sensitive to increasing nutrient input, and carnivorous plants, which are characteristic of these widespread ecosystem types, may be especially sensitive indicators of N deposition. Botanical carnivory is thought to have evolved in nutrient-poor and well-lit habitats such as bogs because the marginal benefits accruing from carnivory exceed the marginal photosynthetic costs associated with the maintenance of carnivorous organs. However, the production of carnivorous organs can be a phenotypically plastic trait. The northern pitcher plant, Sarracenia purpurea, produces leaves specialized for prey capture and nutrient uptake (pitchers) and leaves that are more efficient at photosynthesis (phyllodia). We hypothesized that relative allocation to these two types of leaves reflects ambient nitrogen availability. We manipulated nutrient availability to plants with leaf enrichment and whole-plot fertilization experiments. Increased nitrogen, but not phosphorus, reduced production of pitchers relative to phyllodia; this result provided empirical support for the cost-benefit model of the evolution of botanical carnivory. Because this phenotypic shift in leaf production occurs in ecological time, our results suggest that S. purpurea could be a reliable and inexpensive biological indicator of nitrogen deposition rates. This suggestion is supported by field observations across a geographic gradient of nitrogen deposition.

Animals↗
PubMed2007

When trade-offs interact: balance of terror enforces dominance discovery trade-off in a local ant assemblage.

1. Trade-offs underpin local species coexistence. Trade-offs between interference and exploitative competitive ability provie a mechanism for explaining species coexistence within guilds that exploit overlapping resources. 2. Omnivorous, leaf litter ants exploit a shared food base and occur in species-rich assemblages. In these assemblages, species that excel at usurping food items from other species are poor at finding food items first. In assemblages where some members are attacked by phorid fly parasitoids, host species face an additional trade-off between defending themselves against parasitic attack and maximizing their competitive abilities. Host species thus face two trade-offs that interact via the trait-mediated indirect interaction generated by phorid defence behaviour. 3. In this study we test for the existence of these trade-offs and evaluate the predictions of a model for how they interact in an assemblage of woodland ants in which two behaviourally dominant members are attacked by phorid fly parasitoids as they attempt to harvest food resources. 4. The major findings are that unparasitized species in the assemblage follow a dominance-discovery trade-off curve. When not subject to attack by phorid flies, host species violate that trade-off by finding resources too quickly for their level of behavioural dominance. In contrast, when attacked by their phorid parasitoids, the host species dominance drops such that they fall into the assemblage trade-off. 5. These results match the predictions of the balance of terror model, which derives the optimal host response to parasitism, indicating that the host species balance the competing fitness costs of reduced competitive dominance and loss of workers to parasitism. This result supports the view that understanding the structure of ecological communities requires incorporating the indirect effects created by trait plasticity.

Animals↗
PubMed2006

Linking physiological processes with mangrove forest structure: phosphorus deficiency limits canopy development, hydraulic conductivity and photosynthetic carbon gain in dwarf Rhizophora mangle.

Spatial gradients in mangrove tree height in barrier islands of Belize are associated with nutrient deficiency and sustained flooding in the absence of a salinity gradient. While nutrient deficiency is likely to affect many parameters, here we show that addition of phosphorus (P) to dwarf mangroves stimulated increases in diameters of xylem vessels, area of conductive xylem tissue and leaf area index (LAI) of the canopy. These changes in structure were consistent with related changes in function, as addition of P also increased hydraulic conductivity (Ks), stomatal conductance and photosynthetic assimilation rates to the same levels measured in taller trees fringing the seaward margin of the mangrove. Increased xylem vessel size and corresponding enhancements in stem hydraulic conductivity in P fertilized dwarf trees came at the cost of enhanced mid-day loss of hydraulic conductivity and was associated with decreased assimilation rates in the afternoon. Analysis of trait plasticity identifies hydraulic properties of trees as more plastic than those of leaf structural and physiological characteristics, implying that hydraulic properties are key in controlling growth in mangroves. Alleviation of P deficiency, which released trees from hydraulic limitations, reduced the structural and functional distinctions between dwarf and taller fringing tree forms of Rhizophora mangle.

Phosphorus↗
PubMed2005

Costly carotenoids: a trade-off between predation and infection risk?

Carotenoid reserves in copepods seem costly in terms of predation risk because they make individuals conspicuous. However, carotenoids also seem to play an important role in immune defence as free radical scavengers. To test whether predation risk influences carotenoid levels and whether changes in carotenoid levels are related to changes in immune defence, I examined individual changes in large carotenoid and other lipid droplets upon exposure to predation risk and subsequent exposure to parasites in the copepod Macrocyclops albidus. Copepods reduced carotenoid reserves upon exposure to predators, through which they potentially avoided the costs of being conspicuous under predation risk. Thus, the size of carotenoid reserves is a plastic trait. Such a decrease in carotenoid reserves may also have a negative impact on the copepods' immune system as individuals that decreased their reserves suffered higher parasite prevalence upon exposure to the cestode Schistocephalus solidus. These results suggest that carotenoid reserves may be individually optimized to trade-off each individual's unique costs (predation risk) and benefits (immune defence) of having these reserves.

Animals↗
PubMed2006

Effects of warming conditions in eastern North American forests on red-backed salamander morphology.

Several studies have reported climate-associated changes in phenotypically plastic traits of amphibians, yet it remains unknown whether amphibians can manifest an evolutionary response to global climate change at the rate and magnitude that it is occurring. To assess this issue, we examined temporal change in the morphology of the red-backed salamander (Plethodon cinereus), a small, abundant woodland salamander distributed widely in eastern North America with two distinct morphotypes: striped individuals associated with cooler microclimates and unstriped individuals associated with warmer microclimates. We compiled morph frequencies for 50,960 individual salamanders from 558 sites as recorded in the published literature and in unpublished field notes of herpetologists between 1908 and 2004. We observed that striping probability increased with increasing latitude, longitude, and elevation and decreased (from 80% to 74% range wide) with time. The combined forces of regional climate warming and, particularly, forest disturbance have evidently been sufficient to cause morphological evolution in this amphibian over the last century.

Animals↗
PubMed2004

Ecotypic differentiation and phenotypic plasticity in reproductive traits of Armadillidium vulgare (Isopoda: Oniscidea).

Armadillidium vulgare differed in growth and survivorship on two field sites. Growth rates were higher at a site with consistently higher quality food than at the other site where less high-quality food was produced and which was less predictably accessible. Survivorship was higher at the second site where temperature fluctuations were consistently smaller. Individuals from the two populations were kept for 6 months under the same food and temperature conditions and patterns of resource allocation to reproductive traits analysed. Members of the population from the site with good growth conditions had significantly higher reproductive allocation, by 13.5%, and larger broods, by 9.1%, than those from the site with poor growth conditions. Contrary to theoretical predictions, they also had significantly larger offspring, by 7.5% dry mass. Larger offspring survived better than small ones. This differential survivorship, by 20% for a 3.4% difference in live mass, was much more pronounced under conditions of moisture stress and under fluctuating temperature regimes. Larger offspring would therefore be at a selective advantage on the site with more severe temperature fluctuations. Phenotypic plasticity in reproductive traits in response to experimental changes in food quality, temperature and crowding were monitored. Reproductive allocation was increased by 20.8% under conditions of higher food quality, by 14.7% at higher temperatures, and by 12.5% under less crowded conditions. Brood size, but not offspring dry mass, increased when food quality increased. When crowding increased by 25.0%, the size of broods remained the same but the dry mass of individual offspring decreased by 11.2%. Members of the population from the site with more variable access to high-quality food showed more plasticity in reproductive traits in response to changes in food supply than members of the population from the site with the more predictable food supply. Members of the population from the site with more stable temperatures showed less plasticity to temperature changes than members of the population from the site with greater temperature fluctuations. It is concluded that the observed microevolutionary processes and phenotypic plasticity have adaptative value as responses to spatial and temporal heterogeneity in environmental conditions.

Adaptation, Physiological↗
PubMed2007

Plasticity in animal personality traits: does prior experience alter the degree of boldness?

Theoreticians predict that animal 'personality' traits may be maladaptive if fixed throughout different contexts, so the present study aimed to test whether these traits are fixed or plastic. Rainbow trout (Onchorhyncus mykiss) were given emboldening or negative experiences in the forms of watching bold or shy individuals responding to novelty or winning or losing fights to examine whether prior experience affected boldness. Bold individuals that lost fights or watched shy demonstrators became more shy by increasing their latency to approach a novel object, whereas shy observers that watched bold demonstrators remained cautious and did not modify their responses to novelty. Shy winners became bolder and decreased their latency to approach a novel object, but shy losers also displayed this shift. In comparison, control groups showed no change in behaviour. Bold fishes given negative experiences reduced their boldness which may be an adaptive response; however, shy fishes may base their strategic decisions upon self-assessment of their relative competitive ability and increase their boldness in situations where getting to resources more quickly ensures they outcompete better competitors.

Adaptation, Biological↗
PubMed2006

Phenotypic plasticity of reproductive traits in response to food availability and photoperiod in white-footed mice (Peromyscus leucopus).

Although most temperate-zone mammals are seasonal breeders, many populations display variation in winter reproductive phenotype. For most mammals, the primary environmental cues regulating reproductive status are food availability and photoperiod, and these two factors can interact in their effect. Low food availability is primarily thought to suppress reproduction by reducing body mass and thereby forcing energy allocations to survival alone. However, because most small mammals rely on an increase in food intake rather than stored nutrients for reproduction, we hypothesized that food availability could act as a signal for low resource availability and affect reproduction even when body condition was not affected. We tested the prediction that restricted food access, without reduced body mass, could alter reproductive responses to short photoperiod. We used genetically distinct lines of white-footed mice (Peromyscus leucopus) derived from a wild population with genetic variation in the neuroendocrine pathway that regulates reproduction in response to environmental cues. The lines were created by artificial selection on gonad size in short photoperiods. Individuals from one line strongly suppress gonadal development in response to short photoperiods, while individuals from the other line suppress gonadal development weakly or not at all. Unresponsive individuals from the selected and an unselected control line were exposed to an intermittent food access protocol that did not affect body mass and only slightly reduced total food intake. We found that restricting food access caused reproductive suppression in short photoperiods but not long photoperiods, with no decrease in body mass. These results provide evidence for an interaction between food and photoperiod that is not dependent upon body condition or energy balance. The results also demonstrate plasticity in the reproductive response to photoperiod of otherwise reproductively nonphotoperiodic white-footed mice.

Analysis of Variance↗
PubMed2004

Epigenetic regulation of translation reveals hidden genetic variation to produce complex traits.

Phenotypic plasticity and the exposure of hidden genetic variation both affect the survival and evolution of new traits, but their contributing molecular mechanisms are largely unknown. A single factor, the yeast prion [PSI(+)], may exert a profound effect on both. [PSI(+)] is a conserved, protein-based genetic element that is formed by a change in the conformation and function of the translation termination factor Sup35p, and is transmitted from mother to progeny. Curing cells of [PSI(+)] alters their survival in different growth conditions and produces a spectrum of phenotypes in different genetic backgrounds. Here we show, by examining three plausible explanations for this phenotypic diversity, that all traits tested involved [PSI(+)]-mediated read-through of nonsense codons. Notably, the phenotypes analysed were genetically complex, and genetic re-assortment frequently converted [PSI(+)]-dependent phenotypes to stable traits that persisted in the absence of [PSI(+)]. Thus, [PSI(+)] provides a temporary survival advantage under diverse conditions, increasing the likelihood that new traits will become fixed by subsequent genetic change. As an epigenetic mechanism that globally affects the relationship between genotype and phenotype, [PSI(+)] expands the conceptual framework for phenotypic plasticity, provides a one-step mechanism for the acquisition of complex traits and affords a route to the genetic assimilation of initially transient epigenetic traits.

Biological Evolution↗
PubMed2005

Indirect consequences of artificial selection on plasticity to light quality in Arabidopsis thaliana.

Covariation between light quality- and photoperiod-mediated phenotypic plasticity was investigated using Arabidopsis thaliana. Three episodes of artificial selection were imposed on an index that quantified the plastic response to reduced red to far-red ratios (R:FR), with higher index values indicating greater plasticity. Relative to control lines, two high plasticity (HP) lines showed 1.6- and 2.4-fold increases in the index; low plasticity (LP) lines showed 1.4- and 1.1-fold decreases. A factorial experiment combining high and low R:FR conditions with long and short photoperiods assessed indirect consequences of selection on plasticity. Despite divergent R:FR-mediated plasticities in HP vs. LP lines, all four lines showed increases in photoperiod-mediated responses and decreases in mean leaf number. Complex relationships among trait means, plasticities and underlying mechanisms caution against generalizing about the genetic architecture of plastic traits. Partially independent developmental and evolutionary responses to R:FR and photoperiod are somewhat unsurprising, given this species' cosmopolitan nature.

Arabidopsis↗
PubMed2000

Phenotypic Integration and the Plasticity of Integration in an Amphicarpic Annual.

In annual plants, including amphicarpic annuals, variation in light availability can evoke phenotypic plasticity in multiple traits. We examined plasticity to light availability of vegetative and reproductive performance traits in Amphicarpaea bracteata (Fabaceae) by developing and evaluating three path-analysis models using data from a greenhouse study. To assess whether light availability altered the phenotypic integration of these performance traits, we examined the models' fit to data collected within a high- or a low-light treatment. We also examined whether a single model or alternate models were required to fit data from the two contrasting light treatments. Using our path-analysis approach, we also made comparisons among three population types: shade- and sun-native populations of the widespread variety A. bracteata var. bracteata and sun-native populations of the variety A. bracteata var. comosa. Although each type of population was somewhat distinctive in patterns of integration and in the plasticity of integration, patterns did not correspond to contrasting ecological affinity, i.e., shade- and sun-native population types or to genetic relatedness of the two varieties. Counter to the prediction that selection regimes involving variation in one or more environmental factor favor intermediate levels of integration, phenotypic integration in A. bracteata was very flexible, with plasticity occurring in a trait-by-trait manner. In particular, there was an inverse relationship between aerial and subterranean reproductive modes in low but not high light. Previous studies of amphicarpic annuals have not observed this type of environment-specific life history trade-off.

Journal Article↗