How is life history variation generated from the genetic resource allocation?

A simple quantitative genetic model is proposed to explain the observed genetic correlation structure of a bruchid beetleCallosobruchus chinensis in terms of two underlying variables: the resource acquisition and the resource allocation. Heritabilities and genetic correlations among age-specific, fecundities are regarded as consequences of genetic variations of the two variables. Genetic correlations are predominantly positive in both predictions and observations. Nonetheless, comparison between observed and predicted values in heritabilities, genetic correlations, and genetic principal components suggested significant genetic variances both of the resource allocation and the resource acquisition. The prediction of the model is discussed in relation, to experimental tests of trade-off in life history evolution.     

Tree,sex and size: Ecological determinants of male vs. female fecundity in three Fagus sylvatica stands

Interindividual variation in fecundities has major consequences on population evolutionary potential, through genetic drift and selection. Using two spatially explicit mating models that analyse the genotypes of seeds and seedlings, we investigated the variation of male and female fecundities within and among three European beech (Fagus sylvatica) stands situated along an altitudinal gradient. Female and male individual fecundity distributions were both skewed in this monoecious species, and we found a higher variance in female as compared to male fecundities. Both female and male fecundities increased with tree size and decreased with density and competition in the neighbourhood, the details of these effects suggesting sex‐specific strategies to deal with the impact of limited resource on fecundity. The studied populations were functionally male‐biased. Among‐individual variations in functional gender were not driven by tree size but by density and competition in the neighbourhood. Femaleness decreased under limited resource availability, an expected consequence of the higher cost of female reproduction. Considering the variation of gene flow and genetic drift across elevation, our results suggest that the adaptive potential could be enhanced by low genetic drift at low elevation, and by high pollen‐mediated gene flow at high elevation. Finally, this study predicts a more efficient response to selection for traits related to male vs. female fitness, for a given selection intensity.     

LABORATORY EVOLUTION OF LIFE-HISTORY TRAITS IN THE BEAN WEEVIL (ACANTHOSCELIDES OBTECTUS): THE EFFECTS OF DENSITY-DEPENDENT AND AGE-SPECIFIC SELECTION

Four types of laboratory populations of the bean weevil (Acanthoscelides obtectus) have been developed to study the effects of density-dependent and age-specific selection. These populations have been selected at high (K) and low larval densities (r) as well as for reproduction early (Y) and late (O) in life. The results presented here suggest that the r- and K-populations (density-dependent selection regimes) have differentiated from each other with respect to the following life-history traits: egg-to-adult viability at high larval density (K > r), preadult developmental time (r > K), body weight (r > K), late fecundity (K > r), total realized fecundity (r > K), and longevity of males (r > K). It was also found that the following traits responded in statistically significant manner in populations subjected to different age-specific selection regimes: egg-to-adult viability (O > Y), body weight (O > Y), early fecundity (Y > O), late fecundity (O > Y), and longevity of females and males (O > Y). Although several life-history traits (viability, body weight, late fecundity) responded in similar manner to both density-dependent and age-specific selection regimes, it appears that underlying genetic and physiological mechanisms responsible for differentiation of the r/K and Y/O populations are different. We have also tested quantitative genetic basis of the bean weevil life-history traits in the populations experiencing density-dependent and age-specific selection. Among the traits traded-off within age-specific selection regimes, only early fecundity showed directional dominance, whereas late fecundity and longevity data indicated additive inheritance. In contrast to age-specific selecton regimes, three life-history traits (developmental time, body size, total fecundity) in the density-sependent regimes exhibited significant dominance effects. Lastly, we have tested the congruence between short-term and long-term effects of larval densities. The comparisons of the outcomes of the r/K selection regimes and those obtained from the low- and high-larval densities revealed that there is no congruence between the selection results and phenotypic plasticity for the analyzed life-history traits in the bean weevil.     

THE EVOLUTIONARY GENETICS OF MALE LIFE-HISTORY CHARACTERS IN DROSOPHILA MELANOGASTER

Alternative models of the maintenance of genetic variability, theories of life-history evolution, and theories of sexual selection and mate choice can be tested by measuring additive and nonadditive genetic variances of components of fitness. A quantitative genetic breeding design was used to produce estimates of genetic variances for male life-history traits in Drosophila melanogaster. Additive genetic covariances and correlations between traits were also estimated. Flies from a large, outbred, laboratory population were assayed for age-specific competitive mating ability, age-specific survivorship, body mass, and fertility. Variance-component analysis then allowed the decomposition of phenotypic variation into components associated with additive genetic, nonadditive genetic, and environmental variability. A comparison of dominance and additive components of genetic variation provides little support for an important role for balancing selection in maintaining genetic variance in this suite of traits. The results provide support for the mutation-accumulation theory, but not the antagonistic-pleiotropy theory of senescence. No evidence is found for the positive genetic correlations between mating success and offspring quality or quantity that are predicted by “good genes” models of sexual selection. Additive genetic coefficients of variation for life-history characters are larger than those for body weight. Finally, this set of male life-history characters exhibits a very low correspondence between estimates of genetic and phenotypic correlations.     

Differences in life history strategies between an invasive and a competing resident seed predator

Competitive interactions may arise from biological invasion if a successful invasive species requires the ecological niche of a resident one. Life-history traits that make a species a successful invader are of particular interest in elucidating both invasion success and how interspecific competition may emerge. In southeastern France, the invasion of cedar forests by the seed chalcid Megastigmus schimitscheki generated competitive relationships with the resident M. pinsapinis for the exploitation of the seed resource. Among the numerous ecological traits allowing these seed predators to exploit their niche successfully, the timing of adult emergence, initial egg load and age-specific realized fecundities of females were investigated to help understanding the issue of such interspecific relationships. Spring adult emergence of M. schimitscheki under natural conditions was significantly earlier than that of M. pinsapinis, suggesting that an advantage for the access to the seed resource for the invasive species may be associated with this trait. Initial egg load was significantly higher in M. schimitscheki than in M. pinsapinis and the analysis of age-specific realized fecundity in semi-natural conditions indicated that both M. schimitscheki and M. pinsapinis females lay a large proportion of their eggs during the early days of their lives. In the light of these findings, both earlier phenology and higher reproduction abilities of M. schimitscheki may have the potential to confer an advantage within a competitive context with M. pinsapinis through enhanced seed resource preemption. This may potentially explain the invasiveness of M. schimitscheki in southern France despite the presence of its closely related competitor M. pinsapinis.     

Fecundity of the invasive marine gastropod Crepidula fornicata near the current northern extreme of its range

The calyptraeid gastropod Crepidula fornicata is native to the eastern coast of the United States but has now become an extremely successful invader along much of the European coastline. As the northern limit of its spread is thought to be determined by an inability of adults to tolerate prolonged exposure to low winter temperatures, this study sought to compare the fecundity of females collected from two sites along the Norwegian coastline with that of females collected from Rhode Island, USA. Few other studies have compared the fecundities of marine invertebrates from invasive populations with those found in native populations. For both populations studied, fecundities increased with increasing shell length. However, contrary to expectations, size‐related fecundities were significantly higher for Norwegian females than for Rhode Island females, with Norwegian females producing larger egg capsules and a greater number of embryos per capsule, but not a greater number of egg capsules per brood. Current evidence suggests that at the northern extreme of its invaded range, the fecundity of C. fornicata is increased rather than compromised.     

Probing the evolution of senescence inDrosophila melanogaster withP-element tagging

Natural populations host a wealth of genetic variation in longevity and age-specific schedules of reproduction. This variation provides critical information for inferring the evolutionary origin of senescence. Patterns of mutational effects on age-specific fecundity and survival provide additional insight to distinguish alternative models of senescence. In this study,P-elements bearing thewhite minigene were inserted at random into a common genetic background, generating lines ofD. melanogaster with single, stable transposon inserts. A series of 48 single-P-element lines revealed statistically significant heterogeneity in both longevity and fecundity. Longevity and early fecundity were only weakly positively correlated (r=0.286,P=0.0398). Both the pooled sample and 30 of the individual lines exhibited a leveling of age-specific mortality at advanced ages, in opposition to the classical demographic models. To the extent that these mutational effects are representative of naturally-occurring mutations in heterogeneous populations, this result presents a problem for the evolutionary theory of senescence. Natural selection is inefficient at removing deleterious mutations that are expressed only at late ages, and selection may not differentiate between mutations whose effects on longevity are post-reproductive. A leveling of the mortality rate would also be seen if mutations whose expression is delayed until very late simply do not occur. A simulation of mutation-selection balance among the 48P-element tagged lines shows that the mean longevity declines monotonically with increasing mutation rate, consistent with the mutation-accumulation model.     

Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

Genetic determinants of weight of fast- and slow-twitch skeletal muscles in old mice

The main goal of the study was to explore the genetic architecture underlying muscle weight in old mice. Weight of soleus, tibialis anterior (TA), extensor digitorum longus (EDL), and gastrocnemius muscles was measured in the C57BL/6J (B6) and DBA/2J (D2) strains and derivative generations: a panel of the BXD recombinant inbred (RI) strains and a B6D2 F₂ intercross at the age of 800 days. The between-strain difference in muscle weight (B6 > D2) ranged between 16% and 38%. Linkage analysis identified suggestive quantitative trait loci (QTL) on Chromosomes (Chr) 2, 6, 7, 8, 19, and X that influenced muscle weight in the 800-day-old group. Comparison of weights at 200, 500, and 800 days revealed a variable effect of age among the four muscles. Linkage analysis in the B6D2 F₂ population combined across the three different age groups identified muscle-, sex-, and age-specific QTL on Chr 1, 2, 3, 5, 6, 8, 9, 11, 13, 17, X, and Y. Genetic factors that influence the rate of weight change (within-strain weight difference at two ages) over the lifespan of BXD RIs were mapped to the markers D2Mit369 and D3Mit130 at the genome-wide p < 0.05 for TA muscle in males (between 200 and 800 days) and females (between 500 and 800 days), respectively. Analysis of all age groups supported previous findings that the genetic effects may be muscle-, age-, and sex-specific.     

Effect of host switching on performance of mass‐reared Trichogramma minutum

The effect of host switching on parasitism by Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) (TM) was examined for parasitoids attacking a factitious host, the Mediterranean flour moth, Ephestia kuehniella Zell. (Lepidoptera: Pyralidae) (MFM), and a natural host, the eastern spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) (SBW). Parasitoids that were switched from rearing on the natural host to rearing on the factitious host had lower realized fecundities on both MFM and SBW eggs than a control line that was maintained continuously on the natural rearing host, SBW eggs. Conversely, T. minutum reared on the factitious host and switched to rearing on the natural host showed greater realized fecundities. This increase in realized fecundity was lost in the first generation after switching parasitoids back to the factitious rearing host. T. minutum that were maintained on the natural rearing host (SBW eggs) showed more variation in realized fecundity and host acceptance of both MFM and SBW eggs, than parasitoids that were reared on MFM eggs.     

The IUD follow-up study: A stochastic model

Recurrence equations are derived for the change in genotype frequencies for general fecundity functions. Statistical models that will result in fecundity functions factorable into male and female contributions are developed. Some attention is given to the genetic structure and equilibrium behavior of single-locus genetic systems with additive fecundities.     

Comparative fecundity and oocyte size of Aristaeomorpha foliacea and Aristeus antennatus in the Greek Ionian Sea (E. Mediterranean) (Decapoda: Aristeidae)

Absolute and relative fecundity of 49 ripe gonads (from the cephalic and the abdominal sections of the same specimen) of Aristaeomorpha foliacea and 71 ripe gonads of Aristeus antennatus caught in the Ionian Sea (E. Mediterranean) and their oocyte size were studied. Both average absolute and relative fecundities estimated for A. antennatus were statistically higher than those estimated for A. foliacea; the average absolute fecundity of A. foliacea was 151 936 oocytes/female and that of A. antennatus was 200 472 oocytes/female and the average relative fecundity of A. foliacea was 5477 oocytes/g and that of A. antennatus was 9386 oocytes/g. Fecundity in both red shrimp species is linearly related to carapace length, body and gonad weight, although, the most precise estimation is provided by the relationship of body weight and fecundity. In the present study, the oocyte size (diameter, volume) of both species could be considered very large. In conclusion, A. foliacea shows lower fecundity than A. antennatus, having a lower ovogenetic activity, but it produces larger oocytes. This is in accordance with the highest value of the gonad weight and gonad volume of A. foliacea exhibited in relation to A. antennatus.     

Quantitative genetics of growth and cryptic evolution of body size in an island population

While evolution occurs when selection acts on a heritable trait, empirical studies of natural systems have frequently reported phenotypic stasis under these conditions. We performed quantitative genetic analyses of weight and hindleg length in a free-living population of Soay sheep (Ovis aries) to test whether genetic constraints can explain previously reported stasis in body size despite evidence for strong positive directional selection. Genetic, maternal and environmental covariance structures were estimated across ontogeny using random regression animal models. Heritability increased with age for weight and hindleg length, though both measures of size were highly heritable across ontogeny. Genetic correlations among ages were generally strong and uniformly positive, and the covariance structures were also highly integrated across ontogeny. Consequently, we found no constraint to the evolution of larger size itself. Rather we expect size at all ages to increase in response to positive selection acting at any age. Consistent with expectation, predicted breeding values for age-specific size traits have increased over a twenty-year period, while maternal performance for offspring size has declined. Re-examination of the phenotypic data confirmed that sheep are not getting larger, but also showed that there are significant negative trends in size at all ages. The genetic evolution is therefore cryptic, with the response to selection presumably being masked at the phenotypic level by a plastic response to changing environmental conditions. Density-dependence, coupled with systematically increasing population size, may contribute to declining body size but is insufficient to completely explain it. Our results demonstrate that an increased understanding of the genetic basis of quantitative traits, and of how plasticity and microevolution can occur simultaneously, is necessary for developing predictive models of phenotypic change in nature.     

Interactions between infant growth and survival: Evidence for selection on age-specific body weight in captive common marmosets (Callithrix jacchus)

The objective of this study is to investigate factors influencing infant survival in captive common marmosets. We investigated the influence of age-specific weight, litter size, caging, and the presence of helpers on survival to 6 months of age in 189 Callithrix jacchus infants. Infant survival was analyzed using Cox Proportional Hazards regression, and fitness functions were plotted to explore the relationship between survival and growth. Results indicate that weights at birth and 120 days significantly affect future survival probability. Litter size significantly influences survival prior to 60 days of age with larger litters having poorer survival. Males and females did not have significantly different survival and the presence of helpers in the group did not influence survival probability. Patterns of survival with respect to age-specific weights suggest stabilizing selection on birth weight and directional selection on weight at 120 days of age. Am. J. Primatol. 42:269–280, 1997. © 1997 Wiley-Liss, Inc.     

The maximum yield problem: Distortion in the yield curve due to age structure

An unselective harvest is sustainable if the per capita removal rate balances the net per capita growth rate of the population. If the function relating the growth rate to population density is known, finding the density at which the maximal total harvest rate may be achieved is a simple exercise in parameter optimization. In age-structured populations the per capita growth rate is related in a complicated way to the age-specific vital rates upon which density feedbacks directly operate. Generally, the function relating the per capita growth rate to density will resemble a log transform of the function relating age-specific fecundities to density. Accordingly, maximum yield is attained at a population density that is closer to the saturation density than we would expect on the basis simply of substituting the functional form of the density dependence of fecundity into the parameter optimization model derived for the case without age structure. The amount of the discrepancy increases with the intensity of the density feedback and with the degree to which the reproduction of a cohort is dominated by reproduction taking place during a span of ages that is small relative to the generation time.     

Comparison of measured and parents' reported height and weight in children and adolescents

The objectives of this study were to (i) compare parent-reported height and weight to measured height and weight in children between ages 2 and 17 years, (ii) investigate correlations between magnitude of error of parent-reported data or refusal to estimate height and weight with gender, race/ethnicity, child's age, and age-specific BMI z-score, and (iii) determine sensitivity and specificity of identifying obese youth based on parent-reported data. The authors studied 1,430 consecutive outpatients between ages 2 and 17 years at an outpatient orthopedic clinic. At the initial visit, parents completed a questionnaire including their child's height and weight; height and weight were then measured. Mean height error was very small, with slight overestimation in boys and underestimation in girls. Mean weight error increased with age (P < 0.001), and girls had a larger mean weight error (1.29 kg, 95% confidence interval (CI): 0.65, 1.45) than boys (0.85 kg, 95% CI: 0.8: 0.58, 1.12). Mean weight error also increased with age-specific BMI z-score (r = 0.32, P < 0.001). Correlation between weight error and age-specific BMI z-score was higher among black children (r = 0.45, P < 0.001) than among Hispanic children (r = 0.37, P < 0.001) and was lowest among white children (r = 0.29, P < 0.001). Refusal or inability to estimate weight did not correlate with age-specific BMI z-score. Twenty-one percent of children who were obese would not be identified by using parent-reported data to calculate the BMI.     

Genotypic interactions in an aphid-host plant relationship: Uroleucon rudbeckiae and Rudbeckia laciniata

Summary Four clones of the aphid Uroleucon rudbeckiae were grown on four clones of the host plant Rudbeckia laciniata. Age-specific fecundities were used to determine the fitness, F _i , of each individual aphid. The analysis of variance for F _i revealed that (1) plant genotype has a significant effect on aphid fitness; (2) there is an interaction between aphid and host plant genotypes with respect to aphid fitness; and (3) aphid fitness is affected by phenotypic differences among individual host plants. Result (2) supports the hypothesis that genotypic interactions between aphid and host may maintain genetic diversity in aphid populations. The results of a preference test, while not significant at customary probability levels, suggested that aphids will choose to feed on plants which confer greater fitness.     

LIFE-HISTORY EVOLUTION IN GUPPIES (POECILIA RETICULATA): 1. PHENOTYPIC AND GENETIC CHANGES IN AN INTRODUCTION EXPERIMENT

Previous investigations (Reznick and Endler, 1982; Reznick, 1982a, 1982b) demonstrated that genetic differences in guppy life histories were associated with differences in predation. Guppies from localities with the pike cichlid Crenicichla alta and associated predators matured earlier, had greater reproductive efforts, and produced more and smaller offspring than did guppies from localities with only Rivulus harti as a potential predator. Crenicichla preys primarily on large, sexually mature size-classes of guppies, while Rivulus preys primarily on small, immature size-classes. These patterns of predation are hypothesized to alter mean age-specific survival. Theoretical treatments of such differences in survival predict the observed trends in age at maturity and reproductive effort. We are using introduction experiments to evaluate the role of predators in selecting for these life-history patterns. The experiment whose results are presented here was conducted in a tributary to the El Cedro River (Trinidad), where a waterfall was the upstream limit to the distribution of all fish except Rivulus. Guppies collected from the Crenicichla locality immediately below the waterfall (the downstream control) were introduced over the waterfall in 1981. This introduction released the guppies from Crenicichla predation, exposed them instead to Rivulus predation only, and also introduced them to a different environment, since the introduction site has greater canopy cover than the site of origin. Changes in guppy life-history patterns can be attributed to predation and/or the environment. Evidence from fish collected and preserved in the field demonstrated that, by mid-1983, guppies from the introduction site above the waterfall matured at larger sizes and produced fewer, larger offspring. There were no consistent differences in reproductive allotment (weight of offspring/total weight). With the exception of reproductive allotment, these patterns are identical to previous comparisons between Rivulus and Crenicichla localities. A laboratory genetics experiment demonstrated that males from the introduction site matured at a later age and at a larger size than did males from the control site downstream, as predicted from the “age-specific predation” hypothesis. No differences between localities were observed for female age and size at maturity or for reproductive effort. The trends for fecundity and offspring size were the reverse of those observed in the field. Because only the males changed in the predicted fashion, it is not possible either to reject or to accept the hypothesis of age-specific predation at this time. We discuss the possible causes for these patterns and the high degree of plasticity in the life history, as evidenced by the differences in fecundity and offspring size between the field and laboratory results.     

Demographic Characteristics of Female Mottled Sculpin, Cottus Bairdi, in the Coweeta Creek Drainage, North Carolina

We quantified: (1) growth rate, (2) length-mass relationships, (3) size- and age-specific fecundity, (4) egg size-frequencies, and (5) size- and age-specific egg diameter relationships for reproductively active female C. bairdi from one of the southern-most extant populations of this species (Coweeta Creek drainage, North Carolina). Gravid females were collected during February and March in 1993–1995, and 1998. Cottus bairdi reached an age of 7+ and 79mm standard length. The youngest and smallest gravid female collected was a 41mm 1+ individual. Mature 1+ females were not uncommon and we collected 21 during our study. All females older than age 2 were mature. Mean fecundity for C. bairdi at Coweeta was 71 eggs (range 9–166 eggs). We found significant positive relationships between fecundity and female length, weight and age. Female length and weight also significantly affected mean egg diameter, although the relationship was not linear. Neither female size or age significantly affected mean maximum egg diameters. Female C. bairdi from the Coweeta Creek drainage possess a unique suite of reproductive characteristics that may represent adaptations to the local selective regime or ecophenotypic variation.     

Mapping quantitative trait loci for murine growth: a closer look at genetic architecture

Over 20 years ago, D. S. Falconer and others launched an important avenue of research into the quantitative of body size growth in mice. This study continues in that tradition by locating quantitative trait loci (QTLs) responsible for murine growth, such as age-specific weights and growth periods, and examining the genetic architecture for body weight. We identified a large number of potential QTLs in an earlier F2 intercross (Intercross I) of the SM/J and LG/J inbred mouse strains. Many of these QTLs are replicated in a second F2 intercross (Intercross II) between the same two strains. These replicated regions provide candidate regions for future fine-mapping studies. We also examined body size and growth QTLs using the combined data set from these two intercrosses, resulting in 96 microsatellite markers being scored for 1045 individuals. An examination of the genetic architecture for age-specific weight and growth periods resulted in locating 20 separate QTLs, which were mainly additive in nature, although dominance was found to affect early growth and body size. QTLs affecting early and late growth were generally distinct, mapping to separate chromosome locations. This QTL pattern indicates largely separate genetic and physiological systems for early and later murine growth, as Falconer suggested. We also found sex-specific QTLs for body size with implications for the evolution of sexual dimorphism.