Determination of Fitness Components of Flies Bearing the Recessive Lethal l(2)M167DTS Mutation with Dominant Heat Sensitivity in Artificial Drosophila melanogaster Populations

Elimination of the heat-sensitive l(2)M167 ^DTS mutation from artificial Drosophila melanogaster populations at constant temperature 25°C and various frequencies of the mutation in the parental generation was studied. Components of fitness of the l(2)M167 ^DTS mutation were estimated in the artificial populations by means of the recurrent model of the dependence of the frequency of this mutation in a given generation on its frequency in the previous generation. The model was solved by a numerical method with limitations on the values of some fitness components obtained in test experiments. According to the limitations and frequencies of the l(2)M167 ^DTS mutation, the leading role and limits of the variation in egg-to-adult viability and female fertility were determined. The previously suggested effect of the positive selection for viability of individuals heterozygous for l(2)M167 ^DTS was confirmed.__________Translated from Genetika, Vol. 41, No. 6, 2005, pp. 767–777.Original Russian Text Copyright © 2005 by Kulikov, Kuznetsov, Marec, Mitrofanov.     

The effect of population density on the elimination dynamics of a recessive lethal mutation l(2)M167DTS from experimental populations of Drosophila melanogaster

Dynamics of the elimination of the temperature-sensitive lethal mutation l(2)M167 ^DTS from experimental populations of Drosophila melanogaster under permissive conditions (25°C) was studied. We have shown a rapid elimination of the mutation from the populations, selection for fitness of heterozygous individuals, association of the selection with high larval density and with the direction of the cross that had produced the founder males of the l(2)M167 ^DTS /+ population, and the effect of relative competitive ability of l(2)M167 ^DTS /+ males on the efficiency of the mutation introduction. Modification systems were shown be involved in fitness selection under conditions of high larval density.Translated from Genetika, Vol. 41, No. 3, 2005, pp. 326–333.Original Russian Text Copyright © 2005 by Kulikov, Marec, Mitrofanov.     

The Effect of Male Mating Competitiveness, Developmental Rate, and Viability of Larvae and Pupae in Drosophila melanogaster Heterozygous for the Temperature-Sensitive Lethal Mutation l(2)M167DTS on the Dynamics of the Mutation Elimination from the Population

The dependence of selection on an introduced mutation l(2)M167 ^DTS on male mating competitiveness, viability, and developmental rate of larvae and pupae of Drosophila melanogaster, heterozygous for this mutation, was examined in population experiments with preset conditions. The limitations of fitness parameters of individuals l(2)M167 ^DTS /+ relative to individuals +/+ were estimated according to the conditions of the experiment and phenotypic characteristics of the mutation studied. Under conditions of limited food supply and dependence on emergence time in each generation, the sequence of female mating was shown to be of less importance than the order of medium utilization by the progeny of a certain genotype related to the male success in the first mating. The limiting factors acting on the l(2)M167 ^DTS mutation were viability and developmental rate.__________Translated from Genetika, Vol. 41, No. 5, 2005, pp. 620–625.Original Russian Text Copyright © 2005 by Kulikov, Marec, Mitrofanov.     

Selection on viability of individuals heterozygous for the temperature-sensitive lethal mutation l(2)M167(DTS) in experimental populations of Drosophila melanogaster

In experiments on introduction of mutation l(2)M167(DTS) in Drosophila melanogaster populations, larval and pupal viability and developmental rate are limiting factors determining the intensity of selection on the l(2)M167(DTS) mutation. Notwithstanding the rapid elimination of the mutation from the population, positive selection for viability was shown, which increased fitness of the mutation carriers in generations. The fitness component viability was estimated in individuals l(2)M167(DTS)/+; relative to that of wild-type individuals, it varied from 0.1 to 1. Factors affecting this trait in overcrowded populations were found.     

Determination of fitness components of flies bearing the recessive lethal l(2)M167(DTS) mutation with dominant heat sensitivity in artificial Drosophila melanogaster populations

Elimination of the heat-sensitive l(2)M167(DTS) mutation from artificial Drosophila melanogaster populations at constant temperature 25 degrees C and various frequencies of the mutation in the parental generation was studied. Components of fitness of the l(2)M167(DTS) mutation were estimated in the artificial populations by means of the recurrent model of the dependence of the frequency of this mutation in a given generation on its frequency in the previous generation. The model was solved by a numerical method with limitations on the values of some fitness components obtained in test experiments. According to the limitations and frequencies of the l(2)M167(DTS) mutation, the leading role and limits of the variation in egg-to-adult viability and female fertility were determined. The previously suggested effect of the positive selection for viability of individuals heterozygous for l(2)M167(DTS) was confirmed.     

The effect of population density on the elimination dynamics of a recessive lethal mutation from experimental populations

Dynamics of the elimination of the temperature-sensitive lethal mutation l(2)M167DTS from experimental populations of Drosophila melanogaster under permissive conditions (25 degrees C) was studied. We have shown a rapid elimination of the mutation from the populations, selection for fitness of heterozygous individuals, association of the selection with high larval density and with the direction of the cross that had produced the founder males of the l(2)M167DTS/+ population, and the effect of relative competitive ability of l(2)M167DTS/+ males on the efficiency of the mutation introduction. Modification systems were shown be involved in fitness selection under conditions of high larval density.     

Fertility of Drosophila melanogaster females affected by mutation l(2)M167 DTS

We studied the fertility of D. melanogaster females heterozygous for the dominant temperature sensitive mutation l(2)M167 ^DTS , which exerts a recessive lethal effect at 25°C, under the conditions of stable temperature regimes 25, 28, and 29°C and changing regimes 25 → 29°C and 29 → 25°C. It was shown that inhibition of total activity of oogenesis due to a decreased number of functioning ovarioles is one of the mechanisms underlying the decreased fertility of l(2)M167 ^DTS /+ females. Analysis of individual fertility of each female confirmed also the role of sterility as a component of fertility of the females. Sterilization was realized due both to full depletion of functioning ovarioles and disturbed mechanism of laying the mature eggs onto a substrate as a result of violation of the feedback blocking normal ovulation, which led to the breakdown of ovarioles and filling of the abdominal cavity with mature oocytes. A significant polymorphism of heterozygous females by their fertility was observed. The intensity of sterilization and mortality of l(2)M167 ^DTS /+ females sharply increased at an elevated temperature (29°C), especially at the pupal stage.     

The effect of male mating competitiveness, developmental rate, and viability of larvae and pupae in Drosophila melanogaster heterozygous for the temperature-sensitive lethal mutation l(2)M167DTS on the dynamics of the mutation elimination from the population

The dependence of selection on an introduced mutation 1(2)M167DTS on male mating competitiveness, viability, and developmental rate of larvae and pupae of Drosophila melanogaster, heterozygous for this mutation, was examined in population experiments with preset conditions. The limitations of fitness parameters of individuals l(2)M167DTS/+ relative to individuals +/+ were estimated according to the conditions of the experiment and phenotypic characteristics of the mutation studied. Under conditions of limited food supply and dependence on emergence time in each generation, the sequence of female mating was shown to be of less importance than the order of medium utilization by the progeny of a certain genotype related to the male success in the first mating. The limiting factors acting on the l(2)M167DTS mutation were viability and developmental rate.     

Transpositions of mobile dispersed genes in Drosophila melanogaster and fitness of stocks

Summary In situ hybridization with polytene chromosomes was used to demonstrate the transposition of mobile dispersed genes (mdg)-1 and 3 following the selection of flies from low reproductive activity and vability (LA stock) for high reproductive activity, viability and fitness (LA⁺ and HA stocks).The inbred LA stock is continuously selected for low reproductive activity and viability and maintains at least for twentyfive generations a characteristic pattern of mdg-1 distribution in 14–15 sites. Inbred LA⁺ and HA stocks exhibit a changed pattern of mdg-1 locations and the number of sites reaches 21–25. Parallel and independent selection for higher viability may lead to similar characteristic changes in the localization of mdg-1.In several independent experiments we observed, within one generation, a spontaneous and saltatory growth of viability and fitness in the mass-bred LA stock. In these cases new mdg-1 and mdg-3 sites reproducibly appeared to within several bands, some of them characteristic of LA⁺ and HA stocks.We discuss the possible role of mdg in determining the quantitative characters of individuals and their fitness.     

The fatty acid constitution and ordering state of membranes in dominant temperature-sensitive lethal mutation and wild-typeDrosophila melanogaster larvae

The ordering state and changes in fatty acid composition of microsomal (MS) and mitochondrial (MC) membranes of two dominant temperature-sensitive (DTS) lethal mutations and the wild-type Oregon-R strain larvae ofDrosophila melanogaster have been studied at 18 and 29°C and after temperature-shift experiments. The membranes of wild-type larvae have a stable ordering state, with “S” values between 0.6 (18°C) and 0.5 (29°C) in both membranes which remained unchanged in shift experiments, although the ratios of saturated/unsaturated fatty acids were changed as expected. The stronglyDTS mutation1(2)10 ^DTS forms very rigid membranes at the restrictive temperature (29°C) which cannot be normalized after shift down, while shift up or development at the permissive temperature results in normal ordering state. This mutant is less able to adjust MS and MC fatty acid composition in response to the growth temperature than the wild type. The less temperature-sensitive1(2)2 ^DTS allele occupies an intermediate state between Oregon-R and1(2)10 ^DTS in both respects. We assume and the genetical data suggest that the DTS mutant gene product is in competition with the wild-type product, resulting in a membrane structure which is not able to accommodate to the restrictive temperature.     

Diffusion approximations for one-locus multi-allele kin selection,mutation and random drift in group-structured populations: a unifying approach to selection models in population genetics

Diffusion approximations are ascertained from a two-time-scale argument in the case of a group-structured diploid population with scaled viability parameters depending on the individual genotype and the group type at a single multi-allelic locus under recurrent mutation, and applied to the case of random pairwise interactions within groups. The main step consists in proving global and uniform convergence of the distribution of the group types in an infinite population in the absence of selection and mutation, using a coalescent approach. An inclusive fitness formulation with coefficient of relatedness between a focal individual J affecting the reproductive success of an individual I, defined as the expected fraction of genes in I that are identical by descent to one or more genes in J in a neutral infinite population, given that J is allozygous or autozygous, yields the correct selection drift functions. These are analogous to the selection drift functions obtained with pure viability selection in a population with inbreeding. They give the changes of the allele frequencies in an infinite population without mutation that correspond to the replicator equation with fitness matrix expressed as a linear combination of a symmetric matrix for allozygous individuals and a rank-one matrix for autozygous individuals. In the case of no inbreeding, the mean inclusive fitness is a strict Lyapunov function with respect to this deterministic dynamics. Connections are made between dispersal with exact replacement (proportional dispersal), uniform dispersal, and local extinction and recolonization. The timing of dispersal (before or after selection, before or after mating) is shown to have an effect on group competition and the effective population size. In memory of Sam Karlin.     

Sexual selection,germline mutation rate and sperm competition

Background  

An important component of sexual selection arises because females obtain viability benefits for their offspring from their mate choice. Females choosing extra-pair fertilization generally favor males with exaggerated secondary sexual characters, and extra-pair paternity increases the variance in male reproductive success. Furthermore, females are assumed to benefit from 'good genes' from extra-pair sires. How additive genetic variance in such viability genes is maintained despite strong directional selection remains an evolutionary enigma. We propose that sexual selection is associated with elevated mutation rates, changing the balance between mutation and selection, thereby increasing variance in fitness and hence the benefits to be obtained from good genes sexual selection. Two hypotheses may account for such elevated mutation: (1) Increased sperm production associated with sperm competition may increase mutation rate. (2) Mutator alleles increase mutation rates that are revealed by the expression of condition-dependent secondary sexual characters used by choosy females during their mate choice. M Petrie has independently developed the idea that mutator alleles may account for the maintenance of genetic variation in viability despite strong directional selection.     

Fitness decline under osmotic stress in Caenorhabditis elegans populations subjected to spontaneous mutation accumulation at varying population sizes

The consequences of mutations for population fitness depends on their individual selection coefficients and the effective population size. An earlier study of Caenorhabditis elegans spontaneous mutation accumulation lines evolved for 409 generations at three population sizes found that N_e  = 1 populations declined significantly in fitness whereas the fitness of larger populations (N_e  = 5, 50) was indistinguishable from the ancestral control under benign conditions. To test if larger MA populations harbor a load of cryptic deleterious mutations that are obscured under benign laboratory conditions, we measured fitness under osmotic stress via exposure to hypersaline conditions. The fitness of N_e  = 1 lines exhibited a further decline under osmotic stress compared to benign conditions. However, the fitness of larger populations remained indistinguishable from that of the ancestral control. The average effects of deleterious mutations in N_e  = 1 lines were estimated to be 22% for productivity and 14% for survivorship, exceeding values previously detected under benign conditions. Our results suggest that fitness decline is due to large effect mutations that are rapidly removed via selection even in small populations, with implications for conservation practices. Genetic stochasticity may not be as potent and immediate a threat to the persistence of small populations as other demographic and environmental stochastic factors.     

Epistatic interactions between ancestral genotype and beneficial mutations shape evolvability in Pseudomonas aeruginosa

The idea that interactions between mutations influence adaptation by driving populations to low and high fitness peaks on adaptive landscapes is deeply ingrained in evolutionary theory. Here, we investigate the impact of epistasis on evolvability by challenging populations of two Pseudomonas aeruginosa clones bearing different initial mutations (in rpoB conferring rifampicin resistance, and the type IV pili gene network) to adaptation to a medium containing l ‐serine as the sole carbon source. Despite being initially indistinguishable in fitness, populations founded by the two ancestral genotypes reached different fitness following 300 generations of evolution. Genome sequencing revealed that the difference could not be explained by acquiring mutations in different targets of selection; the majority of clones from both ancestors converged on one of the following two strategies: (1) acquiring mutations in either PA2449 (gcsR, an l ‐serine‐metabolism RpoN enhancer binding protein) or (2) protease genes. Additionally, populations from both ancestors converged on loss‐of‐function mutations in the type IV pili gene network, either due to ancestral or acquired mutations. No compensatory or reversion mutations were observed in RNA polymerase (RNAP) genes, in spite of the large fitness costs typically associated with mutations in rpoB. Although current theory points to sign epistasis as the dominant constraint on evolvability, these results suggest that the role of magnitude epistasis in constraining evolvability may be underappreciated. The contribution of magnitude epistasis is likely to be greatest under the biologically relevant mutation supply rates that make back mutations probabilistically unlikely.     

Space Use and Habitat Selection by Female Louisiana Black Bears in the Tensas River Basin of Louisiana

Abstract: Studies of space use and habitat selection of endangered species are useful for identifying factors that influence fitness of individuals and viability of populations. However, there is a lack of published information regarding these behaviors for the federally threatened Louisiana black bear (Ursus americanus luteolus). We documented space use and habitat selection for 28 female black bears in 2 subpopulations of the Tensas River Basin population in northeast Louisiana, USA. The Tensas subpopulation inhabits a relatively large (>300-km²) contiguous area of bottomland hardwood forest, whereas the Deltic subpopulation exists mainly in 2 small (<7-km²) forested patches surrounded by an agricultural matrix. Females on Deltic maintained smaller seasonal and annual home ranges than females on Tensas (all P < 0.04), except for females with cubs during spring. On Tensas, females with cubs maintained smaller home ranges than females without cubs during spring (P = 0.01), but we did not detect this difference on Deltic or in other seasons. Females on Tensas and Deltic exhibited differences in habitat selection when establishing home ranges and within home ranges (P < 0.001). Deltic females selected mature bottomland hardwood forests and avoided agricultural habitats at both spatial scales. Tensas females selected a mixture of swamps, mature and regenerating forests, and exhibited variation in selection across scale, season, and reproductive status. We suggest that differences in space use and habitat selection between Tensas and Deltic are at least partially due to habitat differences at the landscape (i.e., amount of forested habitat) and patch (i.e., food availability) scales. Our results contribute to the understanding of factors that influence space use and habitat selection by black bears and provide specific information on habitat types selected by Louisiana black bears to agencies involved in habitat protection and restoration for this threatened subspecies.     

Interspecific competition between Drosophila melanogaster and Drosophila simulans: Effects of adult density on adult viability

Survival (viability) of newly eclosed adults of D. simulans st and D. melanogaster Or R-C, which had excess quantities of dead yeast available throughout life, was measured daily for 7 days in an experiment where adult density (6 levels), species frequency (6 levels) and ³²P tissue content (2 levels) were varied factorially. A separate experiment comparing viability in different types of experimental unit also was done. Similar experiments were done for D. simulans st competing against D. melanogaster y w. The entire data were subjected to a least-squares (unbalanced, missing-plot) analysis of variance. D. melanogaster Or R-C had a higher average viability than D. melanogaster y w or D. simulans st, which were equal. The competing strain of D. melanogaster influenced D. simulans st viability — viability being higher when y w was the competitor. Viability decreased over the seven day period but at different rates for the three strains. Increasing density reduced viability for all three strains, but species frequency effects, although significant, were generally not consistent. Females had higher viability than males in both D. melanogaster strains, but the reverse was true in D. simulans st. ³²P lowered viability and experimental unit type altered viability. Numerous interactions were significant.Adult density was shown to have a delayed effect on viability — the delay before the appearance of the effect (an increase in death rate) being decreased as density rose. The term variably delayed density dependent has been adopted to describe the fitness component, adult viability. Some high density populations showed a readjustment (a decrease) in their death rate as a reaction to the effect of reduced density caused by high early mortality.     

On the mode of selection in escherichia coli

Selection experiments between a lac ⁺ wild type and a lac ^– mutant of Escherichia coli were conducted using a serial transfer prcedure at two levels of total sugar concentration, i.e., 0.750 g/l and 1.0 g/l. The relative frequencies of both genotypes were assayed every 48 hours, at the time of transfer and the frequency changes were followed for about four weeks. The frequency of lac ^– decreased steadily and it was eventually eliminated from the mixed populations in all replicate cultures. The regression of logit p_t was shown to be not linear but curvilinear, indicating that fitness differentials are frequency-dependent. The selection is also probably density-dependent.     

Invasion of an indigenous Perna perna mussel bed on the south coast of South Africa by an alien mussel Mytilus galloprovincialis and its effect on the associated fauna

The introduced mussel Mytilus galloprovincialis is progressively increasing in abundance along the south coast of South Africa. Quantitative 0.1 m² samples were collected in the mid-zone of an indigenous Perna perna mussel bed in the 1980s prior to the arrival M. galloprovincialis (12) and in the 2000s during the M. galloprovincialis invasion (16). In addition, in situ counts of M. galloprovincialis were done on eight occasions between 1993 and 2005, and in the low- and high-zones on four occasions. In the mid-zone M. galloprovincialis was absent until 1987, its mean densities were low (< 15 individuals/0.1 m²) between 1993 and 1996, but thereafter increased steadily, peaking in 2004 (at 721 individuals/0.1 m²), before declining in 2005 (331 individuals/0.1 m²). The greatest densities of M. galloprovincialis were recorded at the high-zone (1121 individuals/0.1 m²) and the smallest in the low-zone. As M. galloprovincialis numbers increased, there was an associated, but smaller decline in P. perna numbers and the overall density of mussels increased significantly (P < 0.05). No major change was recorded in the size composition of P. perna. The density of associated fauna differed significantly (P < 0.01) between sampling dates with the lowest and highest values being recorded near the ‘beginning’ (2001) and ‘end’ (2005) of the invasion period respectively. These differences were largely due to variations in the density of barnacles, and the toothed barnacle Chthamalus dentatus appeared to be the only associated faunal species that was directly affected by the M. galloprovincialis invasion, experiencing a significantly (P ≤ 0.05), but temporary decline in density and biomass values.     

Local adaptation and ecological differentiation under selection,migration, and drift in Arabidopsis lyrata*

How the balance between selection, migration, and drift influences the evolution of local adaptation has been under intense theoretical scrutiny. Yet, empirical studies that relate estimates of local adaptation to quantification of gene flow and effective population sizes have been rare. Here, we conducted a reciprocal transplant trial, a common garden trial, and a whole‐genome‐based demography analysis to examine these effects among Arabidopsis lyrata populations from two altitudinal gradients in Norway. Demography simulations indicated that populations within the two gradients are connected by gene flow (0.1 < 4N_em < 11) and have small effective population sizes (N_e < 6000), suggesting that both migration and drift can counteract local selection. However, the three‐year field experiments showed evidence of local adaptation at the level of hierarchical multiyear fitness, attesting to the strength of differential selection. In the lowland habitat, local superiority was associated with greater fecundity, while viability accounted for fitness differences in the alpine habitat. We also demonstrate that flowering time differentiation has contributed to adaptive divergence between these locally adapted populations. Our results show that despite the estimated potential of gene flow and drift to hinder differentiation, selection among these A. lyrata populations has resulted in local adaptation.     

CROSS ABILITY OF MIMULUS GUTTATUS IN RELATION TO COMPONENTS OF GENE FIXATION

The joint effects of parental gene fixation and consanguinity of mates upon the fitness of matings was examined in Mimulus guttatus. Plants from four populations were crossed at five levels of genetic relatedness, and five viability characters were scored in progeny. Parental gene fixation at 12 polymorphic allozyme loci was partitioned into local, subpopulation, and population components. A model is proposed wherein parental gene fixation influences distance-dependent crossing success. At a fixed locus, inbreeding is favored if natural selection caused allele fixation, or is disfavored if gene fixation was random. The distance between mates required to eliminate gene fixation depends upon patch size of fixation. When selective fixation and patch size differ among loci, an optimal crossing distance is possible. In M. guttatus, progeny viability generally decreased with greater relatedness between mates, but this decrease was often heterogeneous among populations. The highest pollen viability and the lowest seed set were found at an intermediate relatedness between mates. To determine whether parental gene fixation influences these crossing patterns, a type of mutational-load analysis was performed. Progeny fitness was regressed on parent F and fitness estimated at F = 1. This was done for each component of F, for a) crosses that maintain gene fixation and b) crosses that eliminate gene fixation. A multiplicative, composite measure of fitness indicates that, in M. guttatus, genes fixed during local or population differentiation favor outbreeding, while genes fixed during subpopulation differentiation favor inbreeding. This predicts that random mating within subpopulations confers highest progeny fitness, exclusive of between-population matings. However, predictions did not fit the observed patterns of crossing success very well, perhaps because gene fixation was relatively low or was not adequately measured at loci influencing fitness.