Interference among deleterious mutations favours sex and recombination in finite populations

Sex and recombination are widespread, but explaining these phenomena has been one of the most difficult problems in evolutionary biology. Recombination is advantageous when different individuals in a population carry different advantageous alleles. By bringing together advantageous alleles onto the same chromosome, recombination speeds up the process of adaptation and opposes the fixation of harmful mutations by means of Muller's ratchet. Nevertheless, adaptive substitutions favour sex and recombination only if the rate of adaptive mutation is high, and Muller's ratchet operates only in small or asexual populations. Here, by tracking the fate of modifier alleles that alter the frequency of sex and recombination, we show that background selection against deleterious mutant alleles provides a stochastic advantage to sex and recombination that increases with population size. The advantage arises because, with low levels of recombination, selection at other loci severely reduces the effective population size and genetic variance in fitness at a focal locus (the Hill-Robertson effect), making a population less able to respond to selection and to rid itself of deleterious mutations. Sex and recombination reveal the hidden genetic variance in fitness by combining chromosomes of intermediate fitness to create chromosomes that are relatively free of (or are loaded with) deleterious mutations. This increase in genetic variance within finite populations improves the response to selection and generates a substantial advantage to sex and recombination that is fairly insensitive to the form of epistatic interactions between deleterious alleles. The mechanism supported by our results offers a robust and broadly applicable explanation for the evolutionary advantage of recombination and can explain the spread of costly sex.     

Sexual selection and the maintenance of sexual reproduction

The maintenance of sexual reproduction is a problem in evolutionary theory because, all else being equal, asexual populations have a twofold fitness advantage over their sexual counterparts and should rapidly outnumber a sexual population because every individual has the potential to reproduce. The twofold cost of sex exists because of anisogamy or gamete dimorphism-egg-producing females make a larger contribution to the zygote compared with the small contribution made by the sperm of males, but both males and females contribute 50% of the genes. Anisogamy also generates the conditions for sexual selection, a powerful evolutionary force that does not exist in asexual populations. The continued prevalence of sexual reproduction indicates that the 'all else being equal' assumption is incorrect. Here I show that sexual selection can mitigate or even eliminate the cost of sex. If sexual selection causes deleterious mutations to be more deleterious in males than females, then deleterious mutations are maintained at lower equilibrium frequency in sexual populations relative to asexual populations. The fitness of sexual females is higher than asexuals because there is no difference in the fecundity of sexual females and asexuals of the same genotype, but the equilibrium frequency of deleterious mutations is lower in sexual populations. The results are not altered by synergistic epistasis in males.     

Higher rates of sex evolve in spatially heterogeneous environments

The evolution and maintenance of sexual reproduction has puzzled biologists for decades. Although this field is rich in hypotheses, experimental evidence is scarce. Some important experiments have demonstrated differences in evolutionary rates between sexual and asexual populations; other experiments have documented evolutionary changes in phenomena related to genetic mixing, such as recombination and selfing. However, direct experiments of the evolution of sex within populations are extremely rare (but see ref. 12). Here we use the rotifer, Brachionus calyciflorus, which is capable of both sexual and asexual reproduction, to test recent theory predicting that there is more opportunity for sex to evolve in spatially heterogeneous environments. Replicated experimental populations of rotifers were maintained in homogeneous environments, composed of either high- or low-quality food habitats, or in heterogeneous environments that consisted of a mix of the two habitats. For populations maintained in either type of homogeneous environment, the rate of sex evolves rapidly towards zero. In contrast, higher rates of sex evolve in populations experiencing spatially heterogeneous environments. The data indicate that the higher level of sex observed under heterogeneity is not due to sex being less costly or selection against sex being less efficient; rather sex is sufficiently advantageous in heterogeneous environments to overwhelm its inherent costs. Counter to some alternative theories for the evolution of sex, there is no evidence that genetic drift plays any part in the evolution of sex in these populations.     

Direct estimation of per nucleotide and genomic deleterious mutation rates in Drosophila

Spontaneous mutations are the source of genetic variation required for evolutionary change, and are therefore important for many aspects of evolutionary biology. For example, the divergence between taxa at neutrally evolving sites in the genome is proportional to the per nucleotide mutation rate, u (ref. 1), and this can be used to date speciation events by assuming a molecular clock. The overall rate of occurrence of deleterious mutations in the genome each generation (U) appears in theories of nucleotide divergence and polymorphism, the evolution of sex and recombination, and the evolutionary consequences of inbreeding. However, estimates of U based on changes in allozymes or DNA sequences and fitness traits are discordant. Here we directly estimate u in Drosophila melanogaster by scanning 20 million bases of DNA from three sets of mutation accumulation lines by using denaturing high-performance liquid chromatography. From 37 mutation events that we detected, we obtained a mean estimate for u of 8.4 x 10(-9) per generation. Moreover, we detected significant heterogeneity in u among the three mutation-accumulation-line genotypes. By multiplying u by an estimate of the fraction of mutations that are deleterious in natural populations of Drosophila, we estimate that U is 1.2 per diploid genome. This high rate suggests that selection against deleterious mutations may have a key role in explaining patterns of genetic variation in the genome, and help to maintain recombination and sexual reproduction.     

The ecological cost of sex

Why sex prevails in nature remains one of the great puzzles of evolution. Sexual reproduction has an immediate cost relative to asexual reproduction, as males only express their contribution to population growth through females. With no males to sustain, an asexual mutant can double its relative representation in the population in successive generations. This is the widely accepted 'twofold cost of males'. Many studies have attempted to explain how sex can recoup this cost from fitness benefits associated with the recombination of parental genotypes, but these require complex biological environments that cycle over evolutionary timescales. In contrast, we have considered the ecological dynamics that govern asexual invasion. Here we show the existence of a threshold growth rate for the sexual population, above which the invasion is halted by intraspecific competition. The asexual population then exerts a weaker inhibitory effect on the carrying capacity of the sexual population than on its own carrying capacity. The stable outcome of this is coexistence on a depleted resource base. Under these ecological circumstances, longer-term benefits of sex may eventually drive out the asexual competitor.     

Sex increases the efficacy of natural selection in experimental yeast populations

Why sex evolved and persists is a problem for evolutionary biology, because sex disrupts favourable gene combinations and requires an expenditure of time and energy. Further, in organisms with unequal-sized gametes, the female transmits her genes at only half the rate of an asexual equivalent (the twofold cost of sex). Many modern theories that provide an explanation for the advantage of sex incorporate an idea originally proposed by Weismann more than 100 years ago: sex allows natural selection to proceed more effectively because it increases genetic variation. Here we test this hypothesis, which still lacks robust empirical support, with the use of experiments on yeast populations. Capitalizing on recent advances in the molecular biology of recombination in yeast, we produced by genetic manipulation strains that differed only in their capacity for sexual reproduction. We show that, as predicted by the theory, sex increases the rate of adaptation to a new harsh environment but has no measurable effect on fitness in a new benign environment where there is little selection.     

青藏高原蒿草属植物繁殖系统生物学特性初探

本文观察和总结了青藏高原蒿草属植物繁育系统的基本特点，并对其特别功能及其进化与适应意义进行了讨论。嵩草属植物具有两个显著的繁殖特点：发达的地下根茎营养繁殖系统和兼性的次级有性恢复系统。各类杂交、突变与染色体各种型式的变化在蒿草属植物中总是同营养繁殖同时存在的；发达的地下根茎营养繁殖系统不仅具有繁殖、传播扩散、占领新生态位的功能，而且还具有使高度适应性的类群或杂种复合类群保持高度纯系性和稳定性，使其优越的遗传基因组合型不至于在有性生殖过程中被破坏或丢失的优势。兼性的次级有性恢复系统使高度适应性和进化性的单纯群在次级遗传生态学过程中转变为有性可育，进一步加强了筛选和甑别适应与进化优秀基因的良性循环。蒿草属植物的繁殖系统蕴藏着极大的居群生态学、遗传生态学和环境适应机制，使蒿草属植物的进化潜力得到了极大的发挥。嵩草属植物具有的这种特殊繁育系统是在对青藏高原环境变化的长期适应中获得的     

Sexual reproduction selects for robustness and negative epistasis in artificial gene networks

The mutational deterministic hypothesis for the origin and maintenance of sexual reproduction posits that sex enhances the ability of natural selection to purge deleterious mutations after recombination brings them together into single genomes. This explanation requires negative epistasis, a type of genetic interaction where mutations are more harmful in combination than expected from their separate effects. The conceptual appeal of the mutational deterministic hypothesis has been offset by our inability to identify the mechanistic and evolutionary bases of negative epistasis. Here we show that negative epistasis can evolve as a consequence of sexual reproduction itself. Using an artificial gene network model, we find that recombination between gene networks imposes selection for genetic robustness, and that negative epistasis evolves as a by-product of this selection. Our results suggest that sexual reproduction selects for conditions that favour its own maintenance, a case of evolution forging its own path.     

Relative fitness can decrease in evolving asexual populations of S. cerevisiae

It is generally accepted from the darwinian theory of evolution that a progressive increase in population adaptation will occur in populations containing genetic variation in fitness, until a stable equilibrium is reached and/or the additive genetic variation is exhausted. However, the theoretical literature of population genetics documents exceptions where mean population fitness may decrease in response to evolutionary changes in gene frequency, due to varying selective coefficients, sexual selection or to epistatic interactions between loci. Until now, no examples of such exceptions have been documented from fitness estimates in either natural or experimental populations. We present here direct evidence that, as a result of epistatic interactions between adaptive mutations, mean population fitness can decrease in asexual evolving populations of the yeast Saccharomyces cerevisiae.     

Sexual reproduction between partners of the same mating type in Cryptococcus neoformans

Cryptococcus neoformans is a globally distributed human fungal pathogen that causes life-threatening meningoencephalitis in immunocompromised patients. It has a defined sexual cycle involving haploid cells of alpha and a mating types, yet the vast majority of environmental and clinical isolates are alpha (ref. 3). Sexual recombination is normally expected to occur between isolates of opposite mating type in organisms with two mating types (or sexes). How sexual reproductive potential can be maintained in an organism with a largely unisexual, nearly clonal population genetic structure is unknown. One clue, however, is that alpha strains undergo fruiting, a process that resembles sexual mating but is thought to be strictly mitotic and asexual. We report here that hallmarks of mating occur during fruiting, including diploidization and meiosis. Pheromone response pathway elements and the key meiotic regulator Dmc1 are required for efficient fruiting. Furthermore, fusion and meiosis can occur between non-isogenic alpha strains, enabling genetic exchange. These studies reveal how sexual reproduction can occur between partners of the same mating type. These findings have implications for the evolution of microbial pathogens, as well as for parthenogenesis, cell fusion events and transitions between self-fertilizing and outcrossing modes of reproduction observed in both fungi and other kingdoms.     

孢子植物的生活史

孢子植物的生活史可归纲为七种类型:(1)只有营养繁殖;(2)只有无性生殖;(3)只有单相植物体进行有性生殖;(4)只有双相植物体进行有性生殖;(5)只有单相植物体进行无性和有性生殖;(6)生活史中出现两种核相的两种植物体;(7)生活史中出现两种核相的三种植物体,后两种均具有世代交替现象。     

北缘普通野生稻实验种群动态研究Ⅱ--生殖生长动态

以江西东乡和湖南茶陵普通野生稻北缘种群为材料,在繁育期对当地构建的普通野生稻种群实验小种群进行了连续的调查和测定,研究北缘普通野生稻在各种实验条件下有性生殖的生长动态,结果表明：(1)不同起始密度下普通野生稻各小种群的有性繁殖量有一定差异,较低的起始密度抽穗率低,而较高起始密度的种群有性繁殖的总量相对较高,到40枝／m^2时抽穗率增长到一个稳定水平．(2)放牧条件对普通野生稻的有性繁殖总量影响不大;而刈割则对普通野生稻的有性繁殖具有明显的抑制作用,在较轻度的切割条件下虽未使普通野生稻个体趋于灭绝,但对抽穗等有性繁殖指标来说则是灾难性的．(3)长期生长于过高的水深对北缘普通野生稻的有性繁殖也具有抑制作用．     

Variation in the reversibility of evolution

How reversible is adaptive evolution? Studies of microbes give mixed answers to this question. Reverse evolution has been little studied in sexual populations, even though the population genetics of sexual populations may be quite different. In the present study, 25 diverged replicated populations of Drosophila melanogaster are returned to a common ancestral environment for 50 generations. Here we show that reverse evolution back to the ancestral state occurs, but is not universal, instead depending on previous evolutionary history and the character studied. Hybrid populations showed no greater tendency to undergo successful reverse evolution, suggesting that insufficient genetic variation was not the factor limiting reverse evolution. Adaptive reverse evolution is a contingent process which occurs with only 50 generations of sexual reproduction.     

Mutation and sex in a competitive world

How do deleterious mutations interact to affect fitness? The answer to this question has substantial implications for a variety of important problems in population biology, including the evolution of sex, the rate of adaptation and the conservation of small populations. Here we analyse a mathematical model of competition for food in which deleterious mutations affect competitive ability. We show that, if individuals usually compete in small groups, then competition can easily lead to a type of genetic interaction known as synergistic epistasis. This means that a deleterious mutation is most damaging in a genome that already has many other deleterious mutations. We also show that competition in small groups can produce a large advantage for sexual populations, both in mean fitness and in ability to resist invasion by asexual lineages. One implication of our findings is that experimental efforts to demonstrate synergistic epistasis may not succeed unless the experiments are redesigned to make them much more naturalistic.     

应用无性繁殖单亲遗传算法实现精馏分离序列优化综合

由于精馏分离序列与二叉树之间具有同构性,在数据结构上精馏分离序列可以抽象为二叉树,直接采用二叉树结构编码方案可以同时表达个体的基因型和表现型．借鉴生物界单亲父本可以经过无性繁殖得到子代的现象,对进化和遗传等重组算子进行设计．应用图论方法建立有效的二叉树结构演化重组机制,从而形成无性繁殖单亲交叉和变异算子．实例表明无性繁殖单亲遗传算法能够成功解算大规模精馏分离序列优化综合问题．     

野古草(Arundinella hirta) 对水淹逆境的生殖响应

分析了嘉陵江边不同强度水淹后的1年生野古草个体的有性生殖和无性生殖数量特征，结果表明：野古草有性生殖植株的单株种子质量、粒数和生殖分配等随水淹强度增大而逐渐降低，高强度水淹后野古草没有进行有性生殖．在一定水淹范围内，表征野古草无性生殖的单株质量、分枝数随水淹强度的增大而呈增大的趋势，说明野古草无性生殖随水淹强度的增加呈增强的趋势．在周期性的自然水淹下，无性生殖是野古草适应自然的生殖方式．     

Fitness of RNA virus decreased by Muller's ratchet

Why sex exists remains an unsolved problem in biology. If mutations are on the average deleterious, a high mutation rate can account for the evolution of sex. One form of this mutational hypothesis is Muller's ratchet. If the mutation rate is high, mutation-free individuals become rare and they can be lost by genetic drift in small populations. In asexual populations, as Muller noted, the loss is irreversible and the load of deleterious mutations increases in a ratchet-like manner with the successive loss of the least-mutated individuals. Sex can be advantageous because it increases the fitness of sexual populations by re-creating mutation-free individuals from mutated individuals and stops (or slows) Muller's ratchet. Although Muller's ratchet is an appealing hypothesis, it has been investigated and documented experimentally in only one group of organisms--ciliated protozoa. I initiated a study to examine the role of Muller's ratchet on the evolution of sex in RNA viruses and report here a significant decrease in fitness due to Muller's ratchet in 20 lineages of the RNA bacteriophage phi 6. These results show that deleterious mutations are generated at a sufficiently high rate to advance Muller's ratchet in an RNA virus and that beneficial, backward and compensatory mutations cannot stop the ratchet in the observed range of fitness decrease.     

淫羊藿无性系种群构件生物量分配格局

淫羊藿为具有克隆生长习性的多年生根茎型草本植物，是重要的药用资源植物．该文以野生淫羊藿为研究材料，于营养生长期和有性生殖期对无性系分株种群进行取样，比较分析不同时期、不同构件生物量分配格局．结果表明，营养期与有性生殖期无性系分株种群生物量差异不显著，但各构件生物量百分比差异显著．营养期根生物量比（RMR）、支持结构生物量比（SBR）和根冠比（R／C）均大于有性生殖期；而叶生物量比（LMR）和叶重分数（LMF）小于有性生殖期．在有性生殖期，营养分株（VR）生物量（83．93％）远远大于生殖分株（RR）生物量（16．07％），用于生殖器官（花）的生物量投资比例非常小（1．45％），叶构件生物量分配比例（50．68％）最大，地下根茎（32．03％）和支持结构（15．84％）生物最分配比例次之．高比例的叶构件生物量有利于合成大量有机物质，这些物质储存于地下根茎中以维持基株在资源缺乏季节的生长，保证种群的生存和繁衍．这些都暗示着淫羊藿不仅有很强的克隆生长能力，而且采取克隆生长为主，有性繁殖为辅的手段维持种群的繁衍与扩散．     

松嫩平原两种群落虎尾草种群有性繁殖的比较

通过调查与测定,分析了松嫩平原两种群落虎尾草种群有性繁殖的特点。结果表明,虎尾草种群的有性繁殖力较强,羊草＋虎尾草群落中虎尾草的生殖株、穗和果实的生长发育均受到种间竞争的制约,籽实生产量明显低于虎尾草群落;不同群落的虎尾草种群生物量分配规律相似,均将较多能量分配给生殖生长,产生大量种子,体现了一年生植物尽可艰保留大量后代这一繁殖策略。     

Clonal reproduction by males and females in the little fire ant

Sexual reproduction can lead to major conflicts between sexes and within genomes. Here we report an extreme case of such conflicts in the little fire ant Wasmannia auropunctata. We found that sterile workers are produced by normal sexual reproduction, whereas daughter queens are invariably clonally produced. Because males usually develop from unfertilized maternal eggs in ants and other haplodiploid species, they normally achieve direct fitness only through diploid female offspring. Hence, although the clonal production of queens increases the queen's relatedness to reproductive daughters, it potentially reduces male reproductive success to zero. In an apparent response to this conflict between sexes, genetic analyses reveal that males reproduce clonally, most likely by eliminating the maternal half of the genome in diploid eggs. As a result, all sons have nuclear genomes identical to those of their father. The obligate clonal production of males and queens from individuals of the same sex effectively results in a complete separation of the male and female gene pools. These findings show that the haplodiploid sex-determination system provides grounds for the evolution of extraordinary genetic systems and new types of sexual conflict.