High temperature intensifies negative density dependence of fitness in red flour beetles

Competition for food, space, or other depletable resources has strong impacts on the fitness of organisms and can lead to a pattern known as negative density dependence, where fitness decreases as population density increases. Yet, many resources that have strong impacts on fitness are nondepletable (e.g., moisture or temperature). How do these nondepletable resources interact with depletable resources to modify negative density dependence? We tested the hypothesis that negative density dependence is modulated by temperature in red flour beetles and tested the prediction that the strength of negative density dependence should decrease as temperature decreases. We measured the number of eggs laid, offspring development time, and the number of offspring that reached maturity at three temperatures and two food treatment combinations as we simultaneously manipulated adult population density. We demonstrated that low temperatures weaken negative density dependence in the number of eggs laid; this pattern was most evident when food was abundant. Density had no effect on development time, but low temperatures increased development time. The percent of eggs that emerged as adults decreased with both density and temperature and increased with food. Temperature, an abiotic driver, can thus modulate density-dependent processes in ectotherms. Therefore, models of population growth for ectotherms should incorporate the effects of temperature.     

Lifetime fitness components in female colour morphs of a damselfly: density‐ or frequency‐dependent selection?

In many damselfly species mature females exhibit colour polymorphism: one female morph resembles the conspecific male (androchrome) while the others do not (gynochromes). Hypotheses for the maintenance of such polymorphisms differ mainly as to whether they are based on density- and/or frequency-dependent selection and on the nature of the frequency dependence. We collected lifetime fitness data (individual lifespan, number of copulations and number of ovipositions) for female morphs of the damselfly Ischnura elegans from 15 insectaries differing in population parameters (density, sex ratio and ratio of andro- to gynochromes). Both density and frequency affected a specific set of the studied fitness components. While morph frequency influenced lifespan, sex ratio influenced the number of copulations, and density affected lifespan and the number of ovipositions. Clearly, discrepancies among studies may be generated if the studied fitness components differ. Our final fitness estimate, the number of ovipositions, was only influenced by density, thereby not supporting frequency-based hypotheses. Contrary to expectation under the current density-based hypothesis, androchromes compared to gynochromes had a lower number of ovipositions at high density. We discuss our findings in the light of mechanisms maintaining the female polymorphism.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 515–523.     

The evolutionary history of Drosophila buzzatii. XVI. Fitness component analysis in an original natural population from Argentina

The adaptive significance of the chromosomal polymorphism of Drosophila buzzati has been studied by means of fitness component analysis in an original population from Argentina. The results show evidence of selection acting through pupal viability, longevity (adult viability) and fecundity on the second chromosome polymorphism, and through pupal viability and virility on the fourth chromosome polymorphism. Changes in chromosomal inversion frequencies throughout the life-cycle suggested an endocyclic pattern of directional selection, which at first seems to be the only detectable mechanism responsible for the maintenance of the polymorphism. However, slow, long-term frequency changes cannot be ruled out. The way in which endocyclic selection acts on this population is different from that in a colonized population previously studied; that is, different fitness components are involved in the maintenance of chromosomal polymorphism. The possible factors that may explain these differences are discussed.     

Intraspecific competition facilitates the evolution of tolerance to insect damage in the perennial plant Solanum carolinense

Tolerance to herbivory (the degree to which plants maintain fitness after damage) is a key component of plant defense, so understanding how natural selection and evolutionary constraints act on tolerance traits is important to general theories of plant–herbivore interactions. These factors may be affected by plant competition, which often interacts with damage to influence trait expression and fitness. However, few studies have manipulated competitor density to examine the evolutionary effects of competition on tolerance. In this study, we tested whether intraspecific competition affects four aspects of the evolution of tolerance to herbivory in the perennial plant Solanum carolinense: phenotypic expression, expression of genetic variation, the adaptive value of tolerance, and costs of tolerance. We manipulated insect damage and intraspecific competition for clonal lines of S. carolinense in a greenhouse experiment, and measured tolerance in terms of sexual and asexual fitness components. Compared to plants growing at low density, plants growing at high density had greater expression of and genetic variation in tolerance, and experienced greater fitness benefits from tolerance when damaged. Tolerance was not costly for plants growing at either density, and only plants growing at low density benefited from tolerance when undamaged, perhaps due to greater intrinsic growth rates of more tolerant genotypes. These results suggest that competition is likely to facilitate the evolution of tolerance in S. carolinense, and perhaps in other plants that regularly experience competition, while spatio-temporal variation in density may maintain genetic variation in tolerance.     

Selfish chromosomal drive shapes recent centromeric histone evolution in monkeyflowers

Centromeres are essential mediators of chromosomal segregation, but both centromeric DNA sequences and associated kinetochore proteins are paradoxically diverse across species. The selfish centromere model explains rapid evolution by both components via an arms-race scenario: centromeric DNA variants drive by distorting chromosomal transmission in female meiosis and attendant fitness costs select on interacting proteins to restore Mendelian inheritance. Although it is clear than centromeres can drive and that drive often carries costs, female meiotic drive has not been directly linked to selection on kinetochore proteins in any natural system. Here, we test the selfish model of centromere evolution in a yellow monkeyflower (Mimulus guttatus) population polymorphic for a costly driving centromere (D). We show that the D haplotype is structurally and genetically distinct and swept to a high stable frequency within the past 1500 years. We use quantitative genetic mapping to demonstrate that context-dependence in the strength of drive (from near-100% D transmission in interspecific hybrids to near-Mendelian in within-population crosses) primarily reflects variable vulnerability of the non-driving competitor chromosomes, but also map an unlinked modifier of drive coincident with kinetochore protein Centromere-specific Histone 3 A (CenH3A). Finally, CenH3A exhibits a recent (<1000 years) selective sweep in our focal population, implicating local interactions with D in ongoing adaptive evolution of this kinetochore protein. Together, our results demonstrate an active co-evolutionary arms race between DNA and protein components of the meiotic machinery in Mimulus, with important consequences for individual fitness and molecular divergence.     

First- and second-order sociality determine survival and reproduction in cooperative cichlids

Cooperative breeders serve as a model to study the evolution of cooperation, where costs and benefits of helping are typically scrutinized at the level of group membership. However, cooperation is often observed in multi-level social organizations involving interactions among individuals at various levels. Here, we argue that a full understanding of the adaptive value of cooperation and the evolution of complex social organization requires identifying the effect of different levels of social organization on direct and indirect fitness components. Our long-term field data show that in the cooperatively breeding, colonial cichlid fish Neolamprologus pulcher, both large group size and high colony density significantly raised group persistence. Neither group size nor density affected survival at the individual level, but they had interactive effects on reproductive output; large group size raised productivity when local population density was low, whereas in contrast, small groups were more productive at high densities. Fitness estimates of individually marked fish revealed indirect fitness benefits associated with staying in large groups. Inclusive fitness, however, was not significantly affected by group size, because the direct fitness component was not increased in larger groups. Together, our findings highlight that the reproductive output of groups may be affected in opposite directions by different levels of sociality, and that complex forms of sociality and costly cooperation may evolve in the absence of large indirect fitness benefits and the influence of kin selection.     

The Effects of Interspecific Competition on the Dynamics of a Polymorphism in an Experimental Population of DROSOPHILA MELANOGASTER

Populations of Drosophila melanogaster with a fourth-chromosome polymorphism were subjected to different levels of competition with Drosophila simulans. The dynamics of the polymorphism and the equilibrium frequencies of the sparkling allele were seen to depend on the competitive level, while the higher productivity of the competing populations was shown to be due to the initial parental density. The effects of competition on fitness components were quantified by fitting the data to both a two-stage selection model and a fertility model. Additional experiments were performed to verify that the interspecific competition caused the changes in fitness. The results are discussed in light of the importance of considering selection components in models of ecological genetics.     

Selection in natural and experimental populations of Drosophila pseudoobscura

The inverted gene arrangements of Drosophila pseudoobscura were used by Th. Dobzhansky in pioneering analyses of natural selection. Recent experiments have shed light on the mechanisms of selection contributing to the balanced polymorphism for the gene arrangements. In experimental populations, both major components of fitness, viability and fertility, are frequency dependent, and rare genotypes often have a selective advantage. Viabilities are also density dependent. The frequency dependence and density dependence of the fitness components are not universal. Some karyotypes are strongly influenced by frequency or density, some are slightly influenced, and some do not appear to be influenced at all. The role of heterozygote advantage in the selection on the gene arrangements is not clear. It is probably one important element in the overall selection, but viability and fertility do not always show a heterozygote advantage. Viability and fertility components of selection seem to be about equally important in changing inversion frequencies. Male mating success is an important component of selection in natural populations, and in one population rare male karyotypes have been found to have a pronounced mating advantage.     

Density-Dependent Fertility Selection in Experimental Populations of DROSOPHILA MELANOGASTER

The effects of larval density on components of fertility fitness were investigated with two mutant lines of Drosophila melanogaster. The differences in adult body weight, wing length, larval survivorship and development time verified that flies reared at high density were resource limited. Experimental results indicate that: (1) relative fecundities of both sexes show density-dependent effects, (2) there is a strong density effect on male and female mating success, and (3) in general, there is a reduction in fecundity differences between genotypes at high density. These results imply that it may be important to consider fertility in models of density-dependent natural selection.     

Variation in the Fitness Effects of Mutations with Population Density and Size in Escherichia coli

The fitness effects of mutations are context specific and depend on both external (e.g., environment) and internal (e.g., cellular stress, genetic background) factors. The influence of population size and density on fitness effects are unknown, despite the central role population size plays in the supply and fixation of mutations. We addressed this issue by comparing the fitness of 92 Keio strains (Escherichia coli K12 single gene knockouts) at comparatively high (1.2×10⁷ CFUs/mL) and low (2.5×10² CFUs/mL) densities, which also differed in population size (high: 1.2×10⁸; low: 1.25×10³). Twenty-eight gene deletions (30%) exhibited a fitness difference, ranging from 5 to 174% (median: 35%), between the high and low densities. Our analyses suggest this variation among gene deletions in fitness responses reflected in part both gene orientation and function, of the gene properties we examined (genomic position, length, orientation, and function). Although we could not determine the relative effects of population density and size, our results suggest fitness effects of mutations vary with these two factors, and this variation is gene-specific. Besides being a mechanism for density-dependent selection (r-K selection), the dependence of fitness effects on population density and size has implications for any population that varies in size over time, including populations undergoing evolutionary rescue, species invasions into novel habitats, and cancer progression and metastasis. Further, combined with recent advances in understanding the roles of other context-specific factors in the fitness effects of mutations, our results will help address theoretical and applied biological questions more realistically.     

Male Aggregation Pheromones Inhibit Ideal Free Habitat Selection in Red Flour Beetles (<Emphasis Type="Italic">Tribolium castaneum</Emphasis>)

The ideal free distribution (IFD) predicts that individuals should be distributed between habitats in proportion to habitat suitability such that mean fitness is equal in each habitat. The IFD is useful in studies examining habitat selection, yet its key assumptions are often violated and the expected IFDs are not consistently detected. While the use of aggregation pheromones by insects is expected to evolve in systems that experience positive density dependence (Allee effect), through a series of experiments we test the hypothesis that aggregation pheromones may limit the ability of individuals to achieve an IFD. Using red flour beetles (Tribolium castaneum), we specifically test the prediction that beetles in groups with an equal sex ratio or in the presence of an artificial aggregation pheromone will deviate from an IFD, whereas female-only groups will achieve an IFD. We also test the hypothesis that aggregation pheromones evolved to promote Allee effects by testing the prediction that beetle fitness will show positive density dependence at low densities. Consistent with our first hypothesis, female groups achieved an IFD, while mixed sex groups and females in the presence of an aggregation pheromone in the low food habitat under-matched the IFD. We found no evidence of Allee effects at low density, but we did find evidence of strong negative density dependence. We demonstrate that the use of aggregation pheromones may negatively impact a population’s ability to achieve an IFD and cast doubt on the hypothesis that male aggregation pheromones evolve to promote Allee effects.     

An Experimental Test of the Relationship between Genetic Variation and Environmental Variation in Tribolium Flour Beetles

The hypothesis that a component of genetic variation for polygenic fitness traits is maintained by environmental heterogeneity was tested using an experimental system involving two species of flour beetles, Tribolium castaneum and T. confusum. Replicated populations of each species from a number of environmental treatments were analyzed for various fitness components following almost 60 generations of natural selection. Environmental differences consisted of flours of cereals commonly invaded by natural populations of these insects.—Tests for adaptation to environments were based on experiments in which populations were reared factorially on each flour, such that population treatment x flour interactions could be detected. Measurements were made of survival, growth rate, larval weight, pupal weight, developmental time, fecundity of individuals at low density and fecundity and cannibalism at high density in both fresh and conditioned media.—Flour differences were found to have significant effects on most traits. Evidence for significant genetic variation and significant genotype x environment interaction was also found. However, no evidence could be found to support the hypothesis that genetic variation was maintained by environmental heterogeneity in food resources. The absence of adaptation to the experimental treatments despite the presence of genetic variation in fitness components suggests that pleiotropy may assume an important role in determining net fitness values of polygenes.     

Mutation processes in natural populations of <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> and <Emphasis Type="Italic">Hirundo rustica</Emphasis> from radiation-contaminated regions of Ukraine

Natural populations of Drosophila melanogaster and Hirundo rustica from regions of Ukraine exhibiting different levels of radiation contamination are investigated. Genetic monitoring was performed with respect to such parameters as the frequency of visible sex-linked mutations, frequency of gonad reduction in Drosophila, and rate of interphase manifestations of chromosomal instability in the erythrocytes of the birds. The results attest to a possible opposite dependence of the level of chromosomal instability among swallows and that of the rate of lethal mutations in the sex chromosome of Drosophila on the density of radiation contamination.     

Measurement of Fitness and Its Components in Six Laboratory Strains of DROSOPHILA MELANOGASTER

Six laboratory strains of Drosophila melanogaster were used to measure "net fitness" and its components by interspecific competition with D. hydei using 100 experimental populations. The "total competitive ability," an estimate of net fitness measured in these competition experiments, was tightly correlated with another measure of net fitness, the population size, in single-species experiments (phenotypic correlation r_p = 0.675 and genotypic correlation r_g = 0.997). Other components of fitness were also measured simultaneously, and the correlation with the net fitness was calculated. The very high correlation between two measurements of net fitness and lower correlations between net fitness and components of fitness suggests that these net fitness measures are more reliable estimates of the "real net fitness" than the components of fitness.     

Adaptive nature of chromosomal rearrangement: differential fitness in pocket gophers

A chromosomal centric fusion polymorphism in populations of the plains pocket gopher, Geomys bursarius, was studied to determine the relative fitness associated with the karyotypic phenotypes. There was a greater number of heterozygous individuals than expected χ₁ ²=8.58, P=0.001. Calculations indicate that the viabilities of the two chromosomal homozygotes were only 35 and 76 percent or that of the heterozygote. Differences in fitness values for the chromosomal morphs for Geomys strongly emphasize the possible adaptive nature of the karyotype and provides a primary mechanism for chromosomal evolution, even in species composed of demes of relatively large size. This is the first case of positive chromosomal heterosis in vertebrates. The plains pocket gopher can now be added to the few empirically documented samples of balanced polymorphism.     

Population-Level Density Dependence Influences the Origin and Maintenance of Parental Care

Parental care is a defining feature of animal breeding systems. We now know that both basic life-history characteristics and ecological factors influence the evolution of care. However, relatively little is known about how these factors interact to influence the origin and maintenance of care. Here, we expand upon previous work and explore the relationship between basic life-history characteristics (stage-specific rates of mortality and maturation) and the fitness benefits associated with the origin and the maintenance of parental care for two broad ecological scenarios: the scenario in which egg survival is density dependent and the case in which adult survival is density dependent. Our findings suggest that high offspring need is likely critical in driving the origin, but not the maintenance, of parental care regardless of whether density dependence acts on egg or adult survival. In general, parental care is more likely to result in greater fitness benefits when baseline adult mortality is low if 1) egg survival is density dependent or 2) adult mortality is density dependent and mutant density is relatively high. When density dependence acts on egg mortality, low rates of egg maturation and high egg densities are less likely to lead to strong fitness benefits of care. However, when density dependence acts on adult mortality, high levels of egg maturation and increasing adult densities are less likely to maintain care. Juvenile survival has relatively little, if any, effect on the origin and maintenance of egg-only care. More generally, our results suggest that the evolution of parental care will be influenced by an organism’s entire life history characteristics, the stage at which density dependence acts, and whether care is originating or being maintained.     

Changing proportions of DNA components in Euglena gracilis

The ratios of satellite deoxyribonucleic acid components to chromosomal deoxyribonucleic acid in Euglena gracilis Z were measured by analytical density gradient ultracentrifugation. Chloroplast deoxyribonucleic acid with a buoyant density of 1.685 g/cm³ exhibited a constant ratio to chromosomal deoxyribonucleic acid during exponential growth and increased twofold as the culture reached the end of the exponential growth phase. The quantity of a satellite deoxyribonucleic acid with a buoyant density of 1.691 g/cm³ was not sufficient to measure the ratio to chromosomal deoxyribonucleic acid during exponential growth but increased to approximately equal the quantity of chloroplast deoxyribonucleic acid as the culture approached the end of the exponential growth phase. The quantity of a deoxyribonucleic acid component with a buoyant density of 1.700 g/cm³ was not sufficient to measure the ratio to chromosomal deoxyribonucleic acid during exponential growth but represented approximately one-third of the total deoxyribonucleic acid as the culture entered the stationary phase of growth.     

Density- and Frequency-Dependent Selection at the Mdh-2 Locus in DROSOPHILA PSEUDOOBSCURA

We have studied differences in the number of Drosophila pseudoobscura produced in a culture when the flies differ with respect to two alleles (F and S) at the Mdh-2 locus, which codes for a malate dehydrogenase enzyme. The studies were done at low and at high density in two- and three-genotype combinations (S/S, F/F and S/F), with one-genotype cultures as controls.——Density affects the fitness of the Mdh-2 genotypes. Different genotypes are differently affected, and the genotype of the competitors also makes a difference on the fitness of a given genotype. When three genotypes are present in a culture, particularly at high density, intergenotypic competition is less intense than intragenotypic competition at several frequency combinations. That is, there is "overcompensation": the three genotypes together exploit the environmental resources better than one genotype alone.—The fitness of the genotypes is frequency dependent in both two-genotype and three-genotype combinations. An inverse relationship between frequency and fitness is observed at high density. This may lead to a stable polymorphism, because the fitness of a genotype increases as its frequency decreases.—Forty independent strains, sampled from a natural population, were used in the experiments. This ensures that more than 95% of the variation present in the genome in the natural population is also present is the experimental cultures. It also ensures that the genetic background of the Mdh-2 alleles is randomized in the same way as it is in nature. However, the possibility remains that Mdh-2 alleles in nature are nonrandomly associated with alleles at closely linked loci. If linkage disequilibrium is present in the experiments because it exists in nature, then the observed effects (such as frequency-dependent selection) would affect the Mdh-2 locus in nature as well.     

Modes of Selection Maintaining an Inversion Polymorphism in DROSOPHILA PAULISTORUM

The possibility that fitness relationships associated with an inversion polymorphism in D. paulistorum were frequency dependent was investigated. Using allozymes of tetrazolium oxidase to mark inversions, the effects of genotype frequency, larval density, and culture conditions on fitness were assessed. The proportions of genotypes among egg-laying females were varied, thus changing the expected proportions of progeny produced in the absence of fecundity or viability selection. The genotypes of progeny were determined by electrophoresis and comparisons of the ratio of the numbers of the different genotypes produced to the expected ratio was used to evaluate fitness relationships. Fitness relationships were dependent on genotype frequency, larval density, and culture conditions. Selection was either absent, directional, frequency dependent (favoring rare types), or heterotic depending on density and culture conditions. It is implied that the adaptive value of genetic variants need not be apparent in all environments, or may change with changing conditions. There is evidence for different criteria for selection in the two sexes. These results add to the evidence supporting the importance of frequency-dependent selection. It is argued that for frequency dependence to be of general importance, selection must act on genes in groups, either as an inversion or as lengths of chromosome with integrity maintained by disequilibrium.