Facultative sexual reproduction under frequency-dependent selection on a single locus

The evolution of a facultative sexual strategy that simultaneously produced sexual and asexual individuals was studied theoretically, under negative frequency-dependence of fitness. The organism was considered to be diploid, characterized by two loci concerning fitness and determining sexual strategy, between which a certain degree of linkage existed. The locus concerning fitness was assumed to involve two alleles, resulting in three genotypes, the relative fitness of an individual being defined by a decreasing function of frequency of its own genotype on this locus in the population. The sexual reproductive strategy was considered to be determined by three alleles; asexual, obligate sexual and facultative sexual. Simulations under various linkages between loci and level of frequency dependence of fitness showed that a facultative sexual strategy was generally able to invade and increase in the population. In particular, when the level of frequency dependence was high to some degree, the facultative strain producing many sexual individuals tended to exclusively occupy the population. Namely, the frequency-dependent selection resulted in a predominance of obligate sexual strategy over asexual strategy, simultaneously causing a subordination of the former to the facultative sexual strategy. This indicated that the evolution of sex should be considered carefully with respect to the possibility of invasion of facultative sex.     

The evolution of obligate sex: the roles of sexual selection and recombination

The evolution of sex is one of the greatest mysteries in evolutionary biology. An even greater mystery is the evolution of obligate sex, particularly when competing with facultative sex and not with complete asexuality. Here, we develop a stochastic simulation of an obligate allele invading a facultative population, where males are subject to sexual selection. We identify a range of parameters where sexual selection can contribute to the evolution of obligate sex: Especially when the cost of sex is low, mutation rate is high, and the facultative individuals do not reproduce sexually very often. The advantage of obligate sex becomes larger in the absence of recombination. Surprisingly, obligate sex can take over even when the population has a lower mean fitness as a result. We show that this is due to the high success of obligate males that can compensate the cost of sex.     

The consequences of facultative sex in a prey adapting to predation

A species reproductive mode, along with its associated costs and benefits, can play a significant role in its evolution and survival. Facultative sexuality, being able to reproduce both sexually and asexually, has been deemed evolutionary favourable as the benefits of either mode may be fully realized. In fact, many studies have focused on identifying the benefits of sex and/or the forces selecting for increased rates of sex using facultative sexual species. The costs of either mode, however, can also have a profound impact on a population's evolutionary trajectory. Here, we used experimental evolution and fitness assays to investigate the consequences of facultative sexuality in prey adapting to predation. Specifically, we compared the adaptive response of algal prey populations exposed to constant rotifer predation and which had alternating cycles of asexual and sexual reproduction where sexual episodes were either facultative (sexual and asexual progeny simultaneously propagated) or obligate (only sexual progeny propagated). We found that prey populations with facultative sexual episodes reached a lower final relative fitness and suffered a greater trade‐off in traits under selection, that is defence and competitive ability, as compared to prey populations with obligate sexual episodes. Our results suggest that costs associated with sexual reproduction (germination time) and asexual reproduction (selection interference) were amplified in the facultative sexual prey populations, leading to a reduction in the net advantage of sexuality. Additionally, we found evidence that the cost of sex was reduced in the obligate sexual prey populations because increased selection for sex was observed via the spontaneous production of sexual cells. These results show that certain costs associated with facultative sexuality can affect an organism's evolutionary trajectory.     

The maintenance of heritable variation through social competition

The paradoxical persistence of heritable variation for fitness-related traits is an evolutionary conundrum that remains a preeminent problem in evolutionary biology. Here we describe a simple mechanism in which social competition results in the evolutionary maintenance of heritable variation for fitness related traits. We demonstrate this mechanism using a genetic model with two primary assumptions: the expression of a trait depends upon success in social competition for limited resources; and competitive success of a genotype depends on the genotypes that it competes against. We find that such social competition generates heritable (additive) genetic variation for "competition-dependent" traits. This heritable variation is not eroded by continuous directional selection because, rather than leading to fixation of favored alleles, selection leads instead to allele frequency cycling due to the concerted coevolution of the social environment with the effects of alleles. Our results provide a mechanism for the maintenance of heritable variation in natural populations and suggest an area for research into the importance of competition in the genetic architecture of fitness related traits.     

Age-associated accumulation of 8-hydroxydeoxyguanosine in mitochondrial DNA of human diaphragm

This is the first report that age-associated accumulation of 8-hydroxydeoxyguanosine (8-OH-dG) does occur in human mitochondrial DNA (mtDNA) in muscle of diaphragm. We extracted mtDNA from human diaphragm muscles from differing age groups, and determined the amount of 8-OH-dG by ultramicro-high performance liquid chromatography/mass-spectrometry system. With the same specimen, multiple deletions of mtDNA were detected by electrophoresis after amplification by the polymerase chain reaction method. In subjects below age 55, the level of 8-OH-dG in mtDNA was below 0.02% of the total deoxyguanosine (dG), whereas, in subjects over age 65, the level of 8-OH-dG increased with age at a rate of ca. 0.25% per 10 years, reaching 0.51% at age 85. Moreover, a concomitant increase in multiple deletions was detected with the increase in age. These results suggest that, in younger diaphragms, replication of mtDNA dilutes out 8-OH-dG being not detectable. In the elderly subjects aged over 65, the replication rate might be slowed down leading to the accumulation of 8-OH-dG in mtDNA, which would accelerate the age-associated multiple deletions of mtDNA observed among the subjects.     

Recombination-dependent concatemeric plasmid replication.

The replication of covalently closed circular supercoiled (form I) DNA in prokaryotes is generally controlled at the initiation level by a rate-limiting effector. Once initiated, replication proceeds via one of two possible modes (theta or sigma replication) which do not rely on functions involved in DNA repair and general recombination. Recently, a novel plasmid replication mode, leading to the accumulation of linear multigenome-length plasmid concatemers in both gram-positive and gram-negative bacteria, has been described. Unlike form I DNA replication, an intermediate recombination step is most probably involved in the initiation of concatemeric plasmid DNA replication. On the basis of structural and functional studies, we infer that recombination-dependent plasmid replication shares important features with phage late replication modes and, in several aspects, parallels the synthesis of plasmid concatemers in phage-infected cells. The characterization of the concatemeric plasmid replication mode has allowed new insights into the mechanisms of DNA replication and recombination in prokaryotes.     

AMELIORATION OF THE DELETERIOUS PLEIOTROPIC EFFECTS OF AN ADAPTIVE MUTATION IN BACILLUS SUBTILIS

The deleterious pleiotropic effects of an adaptive mutation may be ameliorated by one of two modes of evolution: (1) by replacement, in which an adaptive mutation with harmful pleiotropic effects is replaced by one that confers an equal benefit but at less cost; or (2) by compensatory evolution, in which natural selection favors modifiers at other loci that compensate for the deleterious effects of the mutant allele. In this study, we have measured the potential of these two modes of evolution to ameliorate the deleterious pleiotropic effects of resistance to the antibiotic rifampicin in the soil bacterium Bacillus subtilis. One approach was to measure the fitness cost of a series of spontaneous rifampicin-resistance mutations from each of several strains. The potential for amelioration by the replacement mode was estimated by the variation in fitness cost among the mutants of a single strain. Another approach was to introduce a series of different rifampicin-resistance alleles into a diversity of strains, and to measure the fitness cost of rifampicin resistance for each allele-by-strain combination. The potential for amelioration by the replacement mode was estimated by the variation in fitness costs among rifampicin-resistance alleles; the potential for compensatory evolution was estimated by variation in the fitness cost of rifampicin resistance among strains. This study has shown that the cost of rifampicin resistance may be ameliorated by both the compensatory and replacement modes.     

Trade-offs between male and female reproduction associated with allozyme variation in phosphoglucoisomerase in an annual plant (Clarkia unguiculata: Onagraceae)

The genotype of an individual for allozymes such as phosphoglucoisomerase (Pgi) is often not neutral with regard to fitness. Studies of several taxa have found consistent fitness differences among Pgi genotypes expressing different allozymes. We conducted a greenhouse experiment with Clarkia unguiculata to determine whether allelic variation at the Pgi-C1 locus may affect components of male and female function. We found significant differences in siring success between pollen donors homozygous for different Pgi alleles. When a mixture of pollen was applied to stigmas under conditions of gametophytic competition (more pollen deposited on stigmas than there are ovules available to fertilize), donors homozygous for the C allele of Pgi sired more seeds per fruit than B-allele donors. Differences between genotypes with respect to female fertility per fruit contrasted with the male advantage associated with the C allele. Recipients homozygous for the C allele produced fruits with more aborted seeds and fewer viable seeds than recipients homozygous for the B allele. These results suggest that allelic variation at a single locus may have opposing effects on male and female reproductive success in C. unguiculata, and that trade-offs between the two types of reproductive success could contribute to the maintenance of variation at the Pgi-C1 locus.     

Deleterious Mutations as an Evolutionary Factor. II. Facultative Apomixis and Selfing

A population with u deleterious mutations per genome per generation is considered in which only those individuals that carry less than a critical number k of mutations are viable. Besides a large number of loci subject to mutation and selection, the genome contains one or two special loci responsible for the mode of reproduction. Amphimixis vs. apomixis and amphimixis vs. selfing are considered separately. In the first case, the genome degradation rate v (= u/square root k) is found to play the decisive role, as in the case of recombination. When v greater than 1.25, obligate amphimixis is established. If v decreases below this value, the alleles with first low and then larger penetrance are fixed, until alleles conferring obligate asexual reproduction become advantageous. The proportion of resources allocated to produce seeds also increases with decrease of v. These results are unlikely to depend on the genetic basis of the mode of reproduction. The result of competition between outcrossing and selfing depends on both u and k, as well as on whether the mutations are recessive. The alleles for selfing with low penetrance are selected against if the mutations are at all recessive. The fitness of alleles with high penetrance depends primarily on u, decreasing when u increases. There may exist conditions when only the alleles providing intermediate selfing rates can be fixed in a population. In other cases a population may exist with either obligate outcrossing or selfing at a high rate. Thus, truncation selection against deleterious mutations may be a factor supporting obligate or facultative sex despite the twofold advantage of apomixis or selfing.     

Two alleles of a developmentally regulated alpha-tubulin locus in Physarum polycephalum replicate on different schedules.

The replication timing of a pair of natural alleles was compared at two alpha-tubulin loci of the Physarum plasmodium. Taking advantage of the naturally synchronous cell cycle of nuclei within the syncytial plasmodium, we analyzed the replication schedule of specific DNA fragments to a resolution of 10-min intervals within a 3-h S phase. At this level of resolution, differences in replication timing between polymorphic alleles at the same locus can be detected in a heterozygote. Specifically, the 3' region of the altA1 allele completes replication at between 20 and 40 min of S phase. The same region of the altA2 allele completes replication at between 40 and 80 min of S phase. In contrast, both alleles at the altB locus replicate concurrently within the first 10 to 15 min of S phase. Previous studies showed that both altA and altB are expressed in the plasmodium, their message levels peaking at mitosis, just minutes before the onset of S phase. However, altB message is detected at substantially higher levels than altA message on Northern (RNA) blots. The temporal windows over which the altA alleles each replicate are very broad in comparison with the levels of mitotic synchrony and altB replication synchrony in a single plasmodium. The allele-specific replication schedule of the altA locus demonstrates that the temporal organization of replicons is not strictly conserved between homologous chromosomes.     

Novel mutations and DNA-based screening in non-Jewish carriers of Tay-Sachs disease.

We have evaluated the feasibility of using PCR-based mutation screening for non-Jewish enzyme-defined carriers identified through Tay-Sachs disease-prevention programs. Although Tay-Sachs mutations are rare in the general population, non-Jewish individuals may be screened as spouses of Jewish carriers or as relatives of probands. In order to define a panel of alleles that might account for the majority of mutations in non-Jewish carriers, we investigated 26 independent alleles from 20 obligate carriers and 3 affected individuals. Eighteen alleles were represented by 12 previously identified mutations, 7 that were newly identified, and 1 that remains unidentified. We then investigated 46 enzyme-defined carrier alleles: 19 were pseudodeficiency alleles, and five mutations accounted for 15 other alleles. An eighth new mutation was detected among enzyme-defined carriers. Eleven alleles remain unidentified, despite the testing for 23 alleles. Some may represent false positives for the enzyme test. Our results indicate that predominant mutations, other than the two pseudodeficiency alleles (739C-->T and 745C-->T) and one disease allele (IVS9+1G-->A), do not occur in the general population. This suggests that it is not possible to define a collection of mutations that could identify an overwhelming majority of the alleles in non-Jews who may require Tay-Sachs carrier screening. We conclude that determination of carrier status by DNA analysis alone is inefficient because of the large proportion of rare alleles. Notwithstanding the possibility of false positives inherent to enzyme screening, this method remains an essential component of carrier screening in non-Jews. DNA screening can be best used as an adjunct to enzyme testing to exclude known HEXA pseudodeficiency alleles, the IVS9+1G-->A disease allele, and other mutations relevant to the subject's genetic heritage.     

Resistance Gene Replacement in the Mosquito Culex pipiens: Fitness Estimation From Long-Term Cline Series

How adaptation appears and is later refined by natural selection has been the object of intense theoretical work. However, the testing of these theories is limited by our ability to estimate the strength of natural selection in nature. Using a long-term cline series, we estimate the selection coefficients acting on different alleles at the same locus to analyze the allele replacement observed in the insecticide resistance gene Ester in the mosquito Culex pipiens in the Montpellier area, southern France. Our method allows us to accurately account for the resistance allele replacement observed in this area since 1986. A first resistance allele appeared early, which was replaced by a second resistance allele providing the same advantage but at a lower cost, itself being replaced by a third resistance allele with both higher advantage and cost. It shows that amelioration of the adaptation (here resistance to insecticide) through allele replacement was successively achieved by selection of first a generalist allele (i.e., with a low fitness variance across environments) and later a specialist allele (i.e., with a large fitness variance across environments). More generally, we discuss how precise estimates of the strength of selection obtained from field data help us understand the process of amelioration of adaptation.     

Environmentally related patterns of reproductive modes in the aphid Myzus persicae and the predominance of two 'superclones' in Victoria, Australia

Asexual organisms that naturally coexist with sexual relatives may hold the key to understanding the maintenance of sex and recombination, a long-standing problem in evolutionary biology. This situation applies to the peach-potato aphid, Myzus persicae, in southeastern Australia where cyclical parthenogens form mixed populations with obligate parthenogens. We collected M. persicae from several areas across Victoria, genotyped them at seven microsatellite loci and experimentally determined their reproductive mode. The geographic distribution of reproductive modes was correlated with two environmental variables that differentially affect obligate and cyclical parthenogens; obligate parthenogens were less frequent in areas with cold winters because they cannot produce frost-resistant eggs while cyclical parthenogens were limited by the availability of their primary host, peach, on which sexual reproduction takes place. Clonal diversity increased with the proportion of cyclical parthenogens in a sample because they tended to have unique microsatellite genotypes, whereas many obligate parthenogens were copies of the same genotype. Two obligately asexual genotypes stood out as being very abundant and widespread, one constituting 24% and the other 17.4% of the entire collection. Both of these highly successful genotypes were present in the majority of all collection sites. Genetic population structure was weak, albeit significant, with a multilocus FST of only 0.021 when samples were reduced to only one representative of each genotype. Interestingly, obligate parthenogens were, on average, more heterozygous and exhibited larger allele size differences between the two alleles at individual loci than cyclical parthenogens. This striking pattern could result from hybridization, for which we have no evidence, or may reflect the previously proposed model of biased mutational divergence of microsatellite alleles within asexual aphid lineages.     

Genetic assimilation in the evolution of bipedalism

The evolution of the bipedal habit and its attendant anatomical specializations constitute the hallmark of our taxonomic family, but aside from numerous adaptive (ultimate cause) scenarios, no discussion of a genetic proximate cause can be found in the literature. On the surface, the evolution of obligate bipedalism involves the inheritance of an acquired character, since the descendants of facultative bipeds who chose to walk bipedally are hypothesized to have become obligate bipeds. This paper attempts to explain the Lamarckian origin of human bipedalism in a neo-Darwinian manner, by recourse to the adaptability of organisms, and the concept of genetic assimilation.     

An inclusive fitness analysis of synergistic interactions in structured populations

We study the evolution of a pair of competing behavioural alleles in a structured population when there are non-additive or ‘synergistic’ fitness effects. Under a form of weak selection and with a simple symmetry condition between a pair of competing alleles, Tarnita et al. provide a surprisingly simple condition for one allele to dominate the other. Their condition can be obtained from an analysis of a corresponding simpler model in which fitness effects are additive. Their result uses an average measure of selective advantage where the average is taken over the long-term—that is, over all possible allele frequencies—and this precludes consideration of any frequency dependence the allelic fitness might exhibit. However, in a considerable body of work with non-additive fitness effects—for example, hawk–dove and prisoner''s dilemma games—frequency dependence plays an essential role in the establishment of conditions for a stable allele-frequency equilibrium. Here, we present a frequency-dependent generalization of their result that provides an expression for allelic fitness at any given allele frequency p. We use an inclusive fitness approach and provide two examples for an infinite structured population. We illustrate our results with an analysis of the hawk–dove game.     

Segregational Drift Constrains the Evolutionary Rate of Prokaryotic Plasmids

Plasmids are extrachromosomal genetic elements in prokaryotes that have been recognized as important drivers of microbial ecology and evolution. Plasmids are found in multiple copies inside their host cell where independent emergence of mutations may lead to intracellular genetic heterogeneity. The intracellular plasmid diversity is thus subject to changes upon cell division. However, the effect of plasmid segregation on plasmid evolution remains understudied. Here, we show that genetic drift during cell division—segregational drift—leads to the rapid extinction of novel plasmid alleles. We established a novel experimental approach to control plasmid allele frequency at the levels of a single cell and the whole population. Following the dynamics of plasmid alleles in an evolution experiment, we find that the mode of plasmid inheritance—random or clustered—is an important determinant of plasmid allele dynamics. Phylogenetic reconstruction of our model plasmid in clinical isolates furthermore reveals a slow evolutionary rate of plasmid-encoded genes in comparison to chromosomal genes. Our study provides empirical evidence that genetic drift in plasmid evolution occurs at multiple levels: the host cell and the population of hosts. Segregational drift has implications for the evolutionary rate heterogeneity of extrachromosomal genetic elements.     

Fitness evolution and the rise of mutator alleles in experimental Escherichia coli populations

We studied the evolution of high mutation rates and the evolution of fitness in three experimental populations of Escherichia coli adapting to a glucose-limited environment. We identified the mutations responsible for the high mutation rates and show that their rate of substitution in all three populations was too rapid to be accounted for simply by genetic drift. In two of the populations, large gains in fitness relative to the ancestor occurred as the mutator alleles rose to fixation, strongly supporting the conclusion that mutator alleles fixed by hitchhiking with beneficial mutations at other loci. In one population, no significant gain in fitness relative to the ancestor occurred in the population as a whole while the mutator allele rose to fixation, but a substantial and significant gain in fitness occurred in the mutator subpopulation as the mutator neared fixation. The spread of the mutator allele from rarity to fixation took >1000 generations in each population. We show that simultaneous adaptive gains in both the mutator and wild-type subpopulations (clonal interference) retarded the mutator fixation in at least one of the populations. We found little evidence that the evolution of high mutation rates accelerated adaptation in these populations.     

The growth advantage in stationary-phase phenotype conferred by rpoS mutations is dependent on the pH and nutrient environment

Escherichia coli cells that are aged in batch culture display an increased fitness referred to as the growth advantage in stationary phase, or GASP, phenotype. A common early adaptation to this culture environment is a mutant rpoS allele, such as rpoS819, that results in attenuated RpoS activity. However, it is important to note that during long-term batch culture, environmental conditions are in flux. To date, most studies of the GASP phenotype have focused on identifying alleles that render an advantage in a specific environment, Luria-Bertani broth (LB) batch culture. To determine what role environmental conditions play in rendering relative fitness advantages to E. coli cells carrying either the wild-type or rpoS819 alleles, we performed competitions under a variety of culture conditions in which either the available nutrients, the pH, or both were manipulated. In LB medium, we found that while the rpoS819 allele confers a strong competitive fitness advantage at basic pH, it confers a reduced advantage under neutral conditions, and it is disadvantageous under acidic conditions. Similar results were found using other media. rpoS819 conferred its greatest advantage in basic minimal medium in which either glucose or Casamino Acids were the sole source of carbon and energy. In acidic medium supplemented with either Casamino Acids or glucose, the wild-type allele conferred a slight advantage. In addition, populations were dynamic under all pH conditions tested, with neither the wild-type nor mutant rpoS alleles sweeping a culture. We also found that the strength of the fitness advantage gained during a 10-day incubation is pH dependent.     

Adaptation in sexuals vs. asexuals: clonal interference and the Fisher-Muller model

Fisher and Muller's theory that recombination speeds adaptation by eliminating competition among beneficial mutations has proved a popular explanation for the advantage of sex. Recent theoretical studies have attempted to quantify the speed of adaptation under the Fisher-Muller model, partly in an attempt to understand the role of "clonal interference" in microbial experimental evolution. We reexamine adaptation in sexuals vs. asexuals, using a model of DNA sequence evolution. In this model, a modest number of sites can mutate to beneficial alleles and the fitness effects of these mutations are unequal. We study (1) transition probabilities to different beneficial mutations; (2) waiting times to the first and the last substitutions of beneficial mutations; and (3) trajectories of mean fitness through time. We find that some of these statistics are surprisingly similar between sexuals and asexuals. These results highlight the importance of the choice of substitution model in assessing the Fisher-Muller advantage of sex.     

抗马拉硫磷淡色库蚊不同基因型的自然内禀增长率及其对抗性演化的影响

本文涉及淡色库蚊(Culex pipiens pallens)抗马拉硫磷纯合子(RR)、杂合子(RS)和敏感纯合子(SS)的自然内禀增长率及其对马拉硫磷抗性演化的影响。RR、RS和SS的内禀增长率分别为0.1118、0.1171和0.1339。RR和RS基因型在无杀虫剂时呈现出繁殖不利性,RR和RS的相对适合度分别是SS的0.65和0.68。 影响淡色库蚊对马拉硫磷进化的某些因子在计算机上进行了模拟。模拟的因子包括R等位基因的起始频率(P_o)、所用马拉硫磷的剂量和迁入率(m)。模拟结果表明(I)不用药时R等位基因衰减是由于RR和RS基因型的繁殖不利性;(2)在起始种群N_o=200,P_o=0.1,使用杀死全部SS和RS的剂量(8ppm)处理,使R等位基因为有效隐性,并且m≥0.15时可阻止马拉硫磷抗性的发生。