THE EVOLUTION OF RESISTANCE TO HERBIVORY IN IPOMOEA PURPUREA. I. ATTEMPTS TO DETECT SELECTION

In this study, we looked for evidence of directional or stabilizing/disruptive selection on plant size and on the level of damage (resistance) caused by four types of herbivores in the annual morning glory Ipomoea purpurea. Selection was estimated by standard phenotypic regression analysis and by regression on breeding values. The phenotypic regression analysis revealed directional selection for all five characters (i.e., plant size and resistance to four types of herbivores) and indicated that plant size and resistance to corn-earworm damage were subject to stabilizing selection. By contrast, the analysis using breeding values revealed directional selection only for plant size and resistance to corn earworms, while none of the characters examined indicated stabilizing or disruptive selection. These results suggest that intermediate levels of damage in I. purpurea are, in general, not maintained by stabilizing selection. Rather, they may reflect either 1) a transient state that exists while directional selection pushes the population toward complete resistance (or, in one case, total absence of resistance) or 2) the evolution of susceptibility to damage by genetic drift.     

Statistical Genetics of an Annual Plant, Impatiens Capensis. II. Natural Selection

Measurement of natural selection on correlated characters provides valuable information on fitness surfaces, patterns of directional, stabilizing, or disruptive selection, mechanisms of fitness variation operating in nature, and possible spatial variation in selective pressures. We examined effects of seed weight, germination date, plant size, early growth, and late growth on individual fitness. Path analysis showed that most characters had direct or indirect effects on individual fitness, indicating directional selection. For most early life-cycle characters, indirect effects via later characters exceed the direct causal effect on fitness. Selection gradients were uniform across the experimental site. There was no evidence for stabilizing or disruptive selection. We discuss several definitions of stabilizing and disruptive selection. Although early events in the life of an individual have important causal effects on subsequent characters and fitness, there is no detectable genetic variance for most of these characters, so little or no genetic response to natural selection is expected.     

Phenotypic selection in natural populations: what limits directional selection?

Studies of phenotypic selection document directional selection in many natural populations. What factors reduce total directional selection and the cumulative evolutionary responses to selection? We combine two data sets for phenotypic selection, representing more than 4,600 distinct estimates of selection from 143 studies, to evaluate the potential roles of fitness trade-offs, indirect (correlated) selection, temporally varying selection, and stabilizing selection for reducing net directional selection and cumulative responses to selection. We detected little evidence that trade-offs among different fitness components reduced total directional selection in most study systems. Comparisons of selection gradients and selection differentials suggest that correlated selection frequently reduced total selection on size but not on other types of traits. The direction of selection on a trait often changes over time in many temporally replicated studies, but these fluctuations have limited impact in reducing cumulative directional selection in most study systems. Analyses of quadratic selection gradients indicated stabilizing selection on body size in at least some studies but provided little evidence that stabilizing selection is more common than disruptive selection for most traits or study systems. Our analyses provide little evidence that fitness trade-offs, correlated selection, or stabilizing selection strongly constrains the directional selection reported for most quantitative traits.     

COMBINING SELECTIVE EPISODES TO ESTIMATE LIFETIME NONLINEAR SELECTION

Studies measuring natural selection acting via different components of fitness may provide insight into such central questions in evolutionary biology as the evolution of life histories and sexual dimorphism. It is often desirable to combine estimates of selection at different episodes to understand how they interact to produce total lifetime selection. When selective episodes are sequential, total directional selection may be calculated by summing directional selection across episodes. However, it is unclear whether lifetime nonlinear (e.g., stabilizing, disruptive, or correlational) selection may be similarly calculated using estimates of quadratic selection from sequential episodes. Here, I show that lifetime quadratic selection depends not only upon the sum total of quadratic selection across episodes but also upon the pattern of directional selection across episodes. In certain cases, the effects of directional selection across episodes may cancel one another, leading to no net directional selection but strong stabilizing selection. This result suggests that true stabilizing selection may be more common than previously thought, especially when the entire life cycle is considered. The equations derived here are easily applicable to empirical data, as is illustrated both with a simulated dataset and with a reanalysis of a study of quadratic selection in dark‐eyed juncos.     

Natural selection on immune defense: A field experiment

Predicting the evolution of phenotypic traits requires an understanding of natural selection on them. Despite its indispensability in the fight against parasites, selection on host immune defense has remained understudied. Theory predicts immune traits to be under stabilizing selection due to associated trade‐offs with other fitness‐related traits. Empirical studies, however, report mainly positive directional selection. This discrepancy could be caused by low phenotypic variation in the examined individuals and/or variation in host resource level that confounds trade‐offs in empirical studies. In a field experiment where we maintained Lymnaea stagnalis snails individually in cages in a lake, we investigated phenotypic selection on two immune defense traits, phenoloxidase (PO)‐like activity and antibacterial activity, in hemolymph. We used a diverse laboratory population and manipulated snail resource level by limiting their food supply. For six weeks, we followed immune activity, growth, and two fitness components, survival and fecundity of snails. We found that PO‐like activity and growth were under stabilizing selection, while antibacterial activity was under positive directional selection. Selection on immune traits was mainly driven by variation in survival. The form of selection on immune defense apparently depends on the particular trait, possibly due to its importance for countering the present parasite community.     

Field estimates of parentage reveal sexually antagonistic selection on body size in a population of Anolis lizards

Sexual dimorphism evolves when selection favors different phenotypic optima between the sexes. Such sexually antagonistic selection creates intralocus sexual conflict when traits are genetically correlated between the sexes and have sex‐specific optima. Brown anoles are highly sexually dimorphic: Males are on average 30% longer than females and 150% heavier in our study population. Viability selection on body size is known to be sexually antagonistic, and directional selection favors large male size whereas stabilizing selection constrains females to remain small. We build on previous studies of viability selection by measuring sexually antagonistic selection using reproductive components of fitness over three generations in a natural population of brown anoles. We estimated the number of offspring produced by an individual that survived to sexual maturity (termed RS^V), a measure of individual fitness that includes aspects of both individual reproductive success and offspring survival. We found directional selection on male body size, consistent with previous studies of viability selection. However, selection on female body size varied among years, and included periods of positive directional selection, quadratic stabilizing selection, and no selection. Selection acts differently in the sexes based on both survival and reproduction and sexual conflict appears to be a persistent force in this species.     

VISUALIZING MULTIVARIATE SELECTION

Recent developments in quantitative-genetic theory have shown that natural selection can be viewed as the multivariate relationship between fitness and phenotype. This relationship can be described by a multidimensional surface depicting fitness as a function of phenotypic traits. We examine the connection between this surface and the coefficients of phenotypic selection that can be estimated by multiple regression and show how the interpretation of multivariate selection can be facilitated through the use of the method of canonical analysis. The results from this analysis can be used to visualize the surface implied by a set of selection coefficients. Such a visualization provides a compact summary of selection coefficients, can aid in the comparison of selection surfaces, and can help generate testable hypotheses as to the adaptive significance of the traits under study. Further, we discuss traditional definitions of directional, stabilizing, and disruptive selection and conclude that selection may be more usefully classified into two general modes, directional and nonlinear selection, with stabilizing and disruptive selection as special cases of nonlinear selection.     

Variation in viability selection among cohorts of juvenile red squirrels (Tamiasciurus hudsonicus)

Abstract.— Selection will result in observable changes in traits only if it acts consistently in space and time, but few estimates of selection in natural populations have been temporally replicated. Here we estimate viability selection on nestling growth rates for 13 cohorts (1989–2001) of red squirrels (Tamiasciurus hudsonicus) from a natural population located in southwestern Yukon, Canada. Directional selection on nestling growth rates varied in magnitude and direction from one cohort to the next. The magnitude of directional selection was relatively weak in most years (median (β= 0.24), but there were episodes of very strong viability selection ((β > 0.5) in some cohorts. We found no evidence of significant stabilizing or disruptive selection on this trait. Examination of viability selection episodes over shorter time periods suggested that the strength of selection on juveniles in this population was positively related to the time scale over which selection was measured. Viability selection from birth to emergence from the natal nest (50 days of age) and from emergence to successful recruitment (100 days of age) were positively correlated, but were both independent of selection on nestling growth rates from recruitment to potential breeding age (one year). The strength of directional selection on growth rates prior to recruitment was negatively correlated with spring temperature whereas selection from recruitment to breeding was positively correlated with the abundance of spruce cones produced in the previous fall. Episodes of strong directional selection from birth to breeding age appear to be due to potentially rare combinations of environmental conditions. As a result, predicting the occurrence of very strong episodes of selection will be extremely difficult, but predicting the microevolutionary responses to observed selection on individual cohorts remains feasible.     

Sexual and lifetime selection on body size in a marine fish: the importance of life-history trade-offs

Many field measurements of viability and sexual selection on body size indicate that large size is favoured. However, life-history theory predicts that body size may be optimized and that patterns of selection may often be stabilizing rather than directional. One reason for this discrepancy may be that field estimates of selection tend to focus on limited components of fitness and may not fully measure life-history trade-offs. We use an 8-year, demographic field study to examine both sexual selection and lifetime selection on body size of a coral reef fish (the bicolour damselfish, Stegastes partitus). Selection via reproductive success of adults was very strong (standardized selection differential=1.04). However, this effect was balanced by trade-offs between large adult size and reduced cumulative survival during the juvenile phase. When we measured lifetime fitness (net reproductive rate), selection was strongly stabilizing and only weakly directional, consistent with predictions from life-history theory.     

LIFE-CYCLE COMPONENTS OF SELECTION IN ERIGERON ANNUUS: I. PHENOTYPIC SELECTION

The magnitude and direction of phenotypic selection on emergence date and seedling size in Erigeron annuus was measured to determine the heterogeneity of selection among sites and the proportion of fitness variance explained by seedling size and emergence date. Three disturbance treatments (open, annual vegetation, perennial vegetation) were imposed to test the hypothesis of stronger selection on seedlings in competitive environments. Selection was most heterogeneous early in the life cycle, with significant spatial heterogeneity in the magnitude of selection on a local scale. The disturbance treatments affected only fecundity selection gradients and selection was strongest in open plots. Significant variation in the sign of selection differentials on emergence date was observed for establishment and fall viability selection episodes; at later stages selection varied in magnitude but not direction. Seedlings in the earliest cohort experienced high mortality during establishment, but increased size and fecundity later in the life cycle. Both stabilizing and disruptive selection on emergence date were observed during establishment, but in general selection was purely directional. At Stony Brook most selection on emergence date operated indirectly through seedling size, whereas at the Weld Preserve direct selection was stronger. There were persistent effects of both seedling emergence date and rosette diameter on adult fitness components, and October rosette diameter explained 18% of the total phenotypic variance in fecundity. Overall, viability fitness components were much more important than fecundity selection. Winter survivorship was the single most important episode of selection.     

The effect of directed and stabilizing selection on the population size of Drosophila melanogaster

Population size as a measure of population fitness was assayed in laboratory lines of Drosophila melanogaster subjected to directional selection for positive photoactivity and stabilizing selection for a complex of metric wing characters. Three experimental designs were used: directional and stabilizing selection (DS); directional selection (D); unselected controls (C). Population size increased under all three experimental designs. The population size of DS and C lines as well as the rate of increase of the DS lines were higher as compared to the D lines. The results suggest that stabilizing selection maintains population fitness at the higher level.     

VARIABLE SELECTION ALONG A SUCCESSIONAL GRADIENT

Patterns of selection were measured in populations of the perennial grass Danthonia spicata from a successional gradient in northern lower Michigan for a five-year period. Phenotypic variation was found both within and among populations for morphological, reproductive, and life-history traits. Two fitness components were measured: fecundity (total number of spikelets produced) and mortality (number of years an individual lived). Multiple-regression analysis of relative reproductive effort (percentage of culms that flowered), culm length, and leaf length against fitness showed substantial variation in the magnitude and direction of selection among the populations and among the fitness components. When three other reproductive traits were added to the analysis, there were no qualitative changes in estimates of directional selection coefficients, but there were pronounced changes in estimates of stabilizing/disruptive selection components. Patterns of selection were concordant with previously measured genetic changes in reproductive effort along the successional gradient but not concordant with genetic changes in culm length and leaf length. These same patterns were found in comparisons of Michigan and North Carolina populations.     

Natural selection on floral traits through male and female function in wild populations of the heterostylous daffodil Narcissus triandrus

Measurements of natural selection in hermaphrodite populations require the analysis of performance through both female and male sex functions. Here, we investigate selection on three floral traits: flower number, flower length, and corona width through both sex functions in natural populations of the tristylous daffodil Narcissus triandrus . Selection through female function was examined in six populations, and in two of these we also estimated male selection gradients using multilocus microsatellite genotyping of parents and offspring. We detected significant directional selection for flower number through female function, and significant stabilizing selection for corona width and flower length through male function. Variation in male reproductive success was strongly influenced by the distance between mates and was significantly higher than variation in female reproductive success in one population, a result consistent with Bateman's principle. However, variation through both sex functions was similar in the other population and there was a significant negative correlation between female and male fitness indicating sex-specific trade-offs in reproductive success. Selection on floral design in N. triandrus was stronger through male than female function probably because floral morphology plays an important role in promoting effective cross-pollen transfer in populations of this heterostylous species.     

LONG-TERM CORRELATED RESPONSE,INTERPOPULATION COVARIATION,AND INTERSPECIFIC ALLOMETRY

A model of long-term correlated evolution of multiple quantitative characters is analyzed, which partitions selection into two components: one stabilizing and the other directional. The model assumes that the stabilizing component is less variable than the directional component among populations. The major result is that, within a population, the responses of characters to selection in the short term differ qualitatively from those in the long term. In the short term, the responses depend on genetic correlations between characters, but in the long term they are only determined by the fitness functions of stabilizing and directional selection, independent of genetic and phenotypic correlations. Treating the stabilizing component as a constant and assuming the directional component to vary among populations, I present formulas for the interpopulation covariation and interspecific allometry, which are functions of the intensity matrix of stabilizing selection. Particular attention is paid to the relationship between intra- and interpopulation correlations.     

EVOLUTION OF VARIATION AND VARIABILITY UNDER FLUCTUATING,STABILIZING, AND DISRUPTIVE SELECTION

How variation and variability (the capacity to vary) may respond to selection remain open questions. Indeed, effects of different selection regimes on variational properties, such as canalization and developmental stability are under debate. We analyzed the patterns of among‐ and within‐individual variation in two wing‐shape characters in populations of Drosophila melanogaster maintained under fluctuating, disruptive, and stabilizing selection for more than 20 generations. Patterns of variation in wing size, which was not a direct target of selection, were also analyzed. Disruptive selection dramatically increased phenotypic variation in the two shape characters, but left phenotypic variation in wing size unaltered. Fluctuating and stabilizing selection consistently decreased phenotypic variation in all traits. In contrast, within‐individual variation, measured by the level of fluctuating asymmetry, increased for all traits under all selection regimes. These results suggest that canalization and developmental stability are evolvable and presumably controlled by different underlying genetic mechanisms, but the evolutionary responses are not consistent with an adaptive response to selection on variation. Selection also affected patterns of directional asymmetry, although inconsistently across traits and treatments.     

Multilevel selection and neighbourhood effects from individual to metapopulation in a wild passerine

Multilevel selection has rarely been studied in the ecological context of animal populations, in which neighbourhood effects range from competition among territorial neighbours to source-sink effects among local populations. By studying a Dupont's lark Chersophilus duponti metapopulation, we analyze neighbourhood effects mediated by song repertoires on fitness components at the individual level (life-span) and population level (growth rate). As a sexual/aggressive signal with strong effects on fitness, birdsong creates an opportunity for group selection via neighbour interactions, but may also have population-wide effects by conveying information on habitat suitability to dispersing individuals. Within populations, we found a disruptive pattern of selection at the individual level and an opposite, stabilizing pattern at the group level. Males singing the most complex songs had the longest life-span, but individuals with the poorest repertoires lived longer than 'average' males, a finding that likely reflects two male strategies with respect to fitness and sexual trait expression. Individuals from groups with intermediate repertoires had the longest life-span, likely benefitting from conspecific signalling to attract females up to the detrimental spread of competitive interactions in groups with superior vocal skills. Within the metapopulation selection was directional but again followed opposite patterns at the two levels: Populations had the highest growth rate when inhabiting local patches with complex repertoires surrounded by patches with simple repertoires. Here the song may impact metapopulation dynamics by guiding prospecting individuals towards populations advertising habitat quality. Two fitness components linked to viability were therefore influenced by the properties of the group, and birdsong was the target of selection, contributing to linking social/sexual processes at the local scale with regional population dynamics.     

MULTIVARIATE ANALYSIS OF SELECTION IN EXPERIMENTAL POPULATIONS DERIVED FROM HYBRIDIZATION OF TWO ECOTYPES OF THE ANNUAL PLANT DIODIA TERES W. (RUBIACEAE)

Morphologically variable F₂ genotypes derived from hybridization of coastal and inland ecotypes of the annual plant Diodia teres were used to identify selection on morphological traits in the natural habitat of each ecotype. These ecotypes occur in very different habitats, and have evolved pronounced morphological differentiation. Selection analysis can suggest whether present patterns of selection can explain morphological differences between ecotypes. F₂ genotypes were characterized morphologically, clonally replicated, and transplanted into the habitat of each ecotype. Selection was measured on six morphological traits. Directional and stabilizing selection occurred on many traits; direction and strength of selection varied sharply at different stages of growth, as revealed by a path-analysis approach that divided selection into a set of independent components. Directional selection favored traits of the native population at the coastal habitat, but less so at the inland habitat. Selection was of sufficient strength to create the observed morphological differences between ecotypes in 25–100 generations, given constant selection and sufficient genetic variation. In effects on fitness, most traits were neither independent nor consistently interactive with other traits. Rather, many traits entered into strong but evanescent interactions affecting particular components of fitness. Observed interactions did not support the hypothesis that the morphology of each ecotype was functionally integrated to a high degree.     

ARE PARALLEL MORPHOLOGIES OF CAVE ORGANISMS THE RESULT OF SIMILAR SELECTION PRESSURES?

The amphipod Gammarus minus is present in both caves and springs, with cave populations showing elaborated (size and antennae) and reduced (eye) characters relative to spring populations. Earlier studies have shown that cave populations resulted from independent invasions of hydrologically isolated subterranean drainages and that there is genetic variation for both elaborated and reduced characters. In this study we tested the hypothesis that a similar pattern of selection on isolated cave populations is responsible for the parallel evolution of cave morphologies. We used variation in mating success and fecundity to test for the presence of directional selection on eye, antennal, and body size characters in a set of cave and spring populations during a series of seasonal cross-sectional samplings. We found significant directional selection for smaller eyes in caves and larger eyes in springs, which supports the hypothesis that selection is responsible for reduced eye size in cave populations. We also found selection for larger body and antennal size in cave populations, which is consistent with the hypothesis that parallel patterns of selection in caves are responsible for the elaboration of body and antennal size. However, we found selection for larger body and antennal size in spring populations that is not consistent with the observed divergence of spring and cave populations. We suggest that unmeasured components of viability selection could be more important in springs than in caves and may act against the selection for larger size found in spring populations.     

Stabilizing selection on nectar concentration in wild Petunia axillaris,as revealed by genetic analysis of pollen dispersal

Most animal-pollinated plants produce nectar as a pollinator reward. Despite the main role that nectar plays in plant-pollinator interactions, the impact of natural variation in nectar traits on realized male fitness is poorly known. Here, we assessed this relation for a wild Petunia axillaris population using paternity-based direct selection gradient analysis, which allowed us also to infer pollen dispersal patterns. Because male fecundity may depend on other traits which could be associated with nectar characteristics (i.e. volume and concentration), we also considered selection on other key reproductive traits. The analysis revealed that P. axillaris was a strict outcrosser, but that successful pollination occurred mainly among neighbours. Individual plants varied greatly in their male fecundity. Nectar concentration, a key feature of nectar that determines its profitability, was subjected to stabilizing selection. Selection through male function also affected corolla area (positive directional selection), corolla tube length (negative directional selection), and floral display size (stabilizing selection), but none of these traits were phenotypically correlated with nectar characteristics. Because nectar concentration affects the ability and foraging efficiency of different flower visitors to feed on nectar, stabilizing selection may reflect either the preference of the most effective pollinators, or antagonistic selection driven by pollinators and non-pollinating nectar consumers.     

ESTIMATING NONLINEAR SELECTION GRADIENTS USING QUADRATIC REGRESSION COEFFICIENTS: DOUBLE OR NOTHING?

The use of regression analysis has been instrumental in allowing evolutionary biologists to estimate the strength and mode of natural selection. Although directional and correlational selection gradients are equal to their corresponding regression coefficients, quadratic regression coefficients must be doubled to estimate stabilizing/disruptive selection gradients. Based on a sample of 33 papers published in Evolution between 2002 and 2007, at least 78% of papers have not doubled quadratic regression coefficients, leading to an appreciable underestimate of the strength of stabilizing and disruptive selection. Proper treatment of quadratic regression coefficients is necessary for estimation of fitness surfaces and contour plots, canonical analysis of the gamma matrix, and modeling the evolution of populations on an adaptive landscape.