A comparison of sexual and asexual replication strategies in a simplified model based on the yeast life cycle

This paper develops simplified mathematical models describing the mutation-selection balance for the asexual and sexual replication pathways in Saccharomyces cerevisiae, or Baker’s yeast. The simplified models are based on the single-fitness-peak approximation in quasispecies theory. We assume diploid genomes consisting of two chromosomes, and we assume that each chromosome is functional if and only if its base sequence is identical to some master sequence. The growth and replication of the yeast cells is modeled as a first-order process, with first-order growth rate constants that are determined by whether a given genome consists of zero, one, or two functional chromosomes. In the asexual pathway, we assume that a given diploid cell divides into two diploids. For the sake of generality, our model allows for mitotic recombination and asymmetric chromosome segregation. In the sexual pathway, we assume that a given diploid cell divides into two diploids, each of which then divide into two haploids. The resulting four haploids enter a haploid pool, where they grow and replicate until they meet another haploid with which to fuse. In the sexual pathway, we consider two mating strategies: (1) a selective strategy, where only haploids with functional chromosomes can fuse with one another; (2) a random strategy, where haploids randomly fuse with one another. When the cost for sex is low, we find that the selective mating strategy leads to the highest mean fitness of the population, when compared to all of the other strategies. When the cost for sex is low, sexual replication with random mating also has a higher mean fitness than asexual replication without mitotic recombination or asymmetric chromosome segregation. We also show that, at low replication fidelities, sexual replication with random mating has a higher mean fitness than asexual replication, as long as the cost for sex is low. If the fitness penalty for having a defective chromosome is sufficiently high and the cost for sex sufficiently low, then at low replication fidelities the random mating strategy has a mean fitness that is a factor of larger than the asexual mean fitness. We argue that for yeast, the selective mating strategy is the one that is closer to reality, which if true suggests that sex may provide a selective advantage under considerably more relaxed conditions than previous research has indicated. The results of this paper also suggest that S. cerevisiae switches from asexual to sexual replication when stressed, because stressful growth conditions provide an opportunity for the yeast to clear out deleterious mutations from their genomes. That being said, our model does not contradict theories for the evolution of sex that argue that sex evolved because it allows a population to more easily adapt to changing conditions.     

Exact Markov chains versus diffusion theory for haploid random mating

Exact discrete Markov chains are applied to the Wright-Fisher model and the Moran model of haploid random mating. Selection and mutations are neglected. At each discrete value of time t there is a given number n of diploid monoecious organisms. The evolution of the population distribution is given in diffusion variables, to compare the two models of random mating with their common diffusion limit. Only the Moran model converges uniformly to the diffusion limit near the boundary. The Wright-Fisher model allows the population size to change with the generations. Diffusion theory tends to under-predict the loss of genetic information when a population enters a bottleneck.     

Pathogenic variation and sexual reproduction in Swedish populations of Bremia lactucae

Summary The host-pathogen interaction between lettuce (Lactuca sativa) and downy mildew (Bremia lactucae) is mainly differential and the resistance so far utilized in the host is vertical. As in many other obligate parasites, the introduction of cultivars with new vertical resistance has exerted a strong selection pressure on the pathogen resulting in significant changes in virulence frequencies and in the establishment of races with new combinations of virulence. Genetic diversity in pathogen populations may arise through mutation and gene flow, and new virulence genotypes may then be established through parasexuality and sexual recombination. In Swedish populations of Bremia lactucae, the pattern of variation in the parasite agrees well with that which might be expected in a diploid, outcrossing organism with frequent sexual reproduction. This is supported by: two or more isolates, different in virulence and mating type, may occur together on the same lettuce leaf; zygotes (oospores) are formed in all populations investigated and the frequency varies from 22% to 98%; oospores germinate rather frequently under suitable conditions. To breed for resistance in dynamic host-pathogen systems such as this one is difficult and the program should preferably be based on race-non-specific resistance.     

The occurrence of sexual reproduction among ant workers

In less than 100 species of ponerine ants, queens no longer exist and have been replaced by mated egg-laying workers. Workers in other subfamilies can lay haploid eggs when queens are removed, but they never reproduce sexually. Ponerine workers are able to mate because they have a spermatheca in most species, foreign males are sexually active near their nests, and their pygidial gland secretions can assume a sexual meaning. Furthermore, ponerine queens are seldom very fecund, and one or several gamergates are able to approximate their egg production. Finally, opportunities for colony fragmentation occur consequent to their life history, and this is a necessary precondition because gamergates cannot start new colonies independently. Many of these characteristics are associated with the limited caste divergence exhibited in this phylogenetically primitive group. Although a few non-ponerine species exhibit some of these preconditions, gamergates have not been found outside the Ponerinae, which alone exhibit the combination of traits leading to queen elimination and worker mating.     

Sexual selection and intelligence: Does sexual reproduction drive the evolution of intelligence?

The basal hypothesis discussed here is the idea that brain architecture could be plastic on a very basal, genetic level due to sexual recombination and reassortment of alleles of genes related to brain development, e.g., neuronal cell adhesion molecules (NCAMs) and others.The role of sexual reassortment leads the study of brain development, species behavior and intelligence to a new version of the so-called “Red Queen Hypothesis”: using the mechanism described here, a kind of runaway selection mechanism seems to arise. Even if NCAMs are almost constant within an individual, they seem to act very differently at the population level and so the role of reassorting polymorphic NCAM- (and other) genes gets particularly clear. If several NCAM-NCAM combinations cause extreme behavior and intelligence variability in a population, these combinations also represent a use of sexual selection. This mechanism of NCAM allele assortment seems to be important for the process of speciation by mutual selection of individuals. Therefore NCAM variants and their associated behaviors are thought to be important for the development of intelligence, in that they promote the attraction of individuals with already high intelligence, leading to the speciation of super-intelligent groups.     

Selection, mutation and sexual reproduction in an infinite haploid population with a genome of finite length

We present a model which describes mutation, selection and sexual reproduction in an infinite haploid population with a finite genome. Each generation is described using an approximation which assures a certain persistent form of the distribution of the number of deleterious elements. The steady state exists and is determined. In addition, we conclude that the introduction of sexual reproduction increases the mean fitness in the equilibrium.     

A selection system for diploid and against haploid cells inSchizosaccharomyces pombe

We have isolated a mutant ofSchizosaccharomyces pombe whose growth is temperature sensitive when it is haploid but not when it is diploid. This mutant may provide a useful system for selecting nonconditional mutants which are defective in diploid formation upon conjugation.     

Trade-offs between sexual and asexual reproduction in the genus Mimulus

Summary Six estimates of resource allocation to sexual reproduction (nectar concentration, nectar volume, sugar production, pollinator visitation rates, fruit-set, and seed production) and a single estimate of resource allocation to asexual reproduction (the rate of rooted-branch production) were examined for five species of Mimulus. There were significant interspecific differences for all paramaters. With the exception of nectar concentration, there was 1) a consistent positive correlation among the parameters measuring allocation to sexual reproduction, and 2) a significant negative correlation between parameters measuring allocation to sexual reproduction and the rate of rooted-branch production (asexual reproduction). The results indicate that Mimulus species which produce the highest volumes of nectar 1) receive the most pollinator visits, 2) have the highest fruit-set, 3) produce the most seeds, and 4) produce the fewest rooted branches.     

Influence of spatial structure on the maintenance of sexual reproduction

We include spatial extension into a model for the maintenance of sexual reproduction introduced recently. The model is based on a broad spectrum of resources, which regrow slowly. Other key features of the model are that sexual reproduction sets in when resources become scarce and that only a few genotypes can coexist locally. The extension of the model to several patches in space is done in two different ways. Model A is based on central egg deposition and allows migration of juveniles into all patches. Model B has a one-dimensional array of patches with migration only between neighboring patches. The main findings are that wide dispersal favors asexuals, while for slower migration there is a wide range of parameter values for which sexually reproducing species always win against asexuals. These results are conform with major patterns for the distribution of parthenogenesis in animals and plants, i.e. the prevalence of parthenogenetic reproduction in minute species, which are easily dispersed by physical forces, such as protists and small metazoans including e.g. bdelloid rotifers, tardigrades and nematodes.     

Estimation of the relative importance of sexual and vegetative reproduction in the clonal woodland herb Anemone nemorosa

Anemone nemorosa is a perennial rhizomatous plant of European woodlands. The “probability of clonal identity” method estimated the relative proportion of sexual to vegetative reproduction in this species to be 4.4% from allozyme genotype distributions. This result is congruent with investigations on the germination, short-term demography, population genetics, and breeding system of this species, and supports the hypothesis that even low levels of seedling recruitment can maintain considerable intrapopulational genetic diversity. Received: 2 March 1998 / Accepted: 24 July 1998     

菊科12种外来植物的有性繁殖特征和入侵风险研究

菊科植物具有极强的有性繁殖能力,对外来菊科观赏植物的引进存在极高的入侵风险。本研究以武汉常见的菊科12种外来观赏植物为对象,通过有性繁殖特征调查,研究植株的花部特征、花粉活力、传粉系统、种子产量,分析各物种有性繁殖能力的差异,评估其入侵能力。结果显示,多数物种可在隔绝传粉者情况下结实;访花昆虫类型多样,共观察到26种访花昆虫,其中膜翅目蜂类是主要的传粉者;结实情况具有一定差异。综合考虑花粉活力、传粉情况和结实情况,认为黑心金光菊(Rudbeckia hirta L.)、松果菊(Echinacea purpurea(L.)Moench)、大花金鸡菊(Coreopsis grandiflora Hogg.)、蛇目菊(Sanvitalia procumbens Lam.)的有性繁殖竞争力较高,对本地传粉环境可能有较大影响。本研究对近缘物种有性繁殖特征的综合分析结果可为评估外来物种的入侵风险提供一定的参考。     

桦纤孔菌有性生殖相关特征及交配型位点分析

对桦纤孔菌菌株MDJCBS88的显微形态、菌丝及担孢子核相进行了观察。采用棉籽壳培养基对担孢子萌发形成的菌株进行栽培试验,筛选出不形成子实体或子实体发育不完整的菌株,将这些菌株在平板上进行了亲和试验,分析桦纤孔菌的有性生殖方式;并基于基因组序列进行交配型基因克隆验证,分析桦纤孔菌的交配型位点结构。显微观察发现,桦纤孔菌菌丝没有锁状联合结构,菌丝细胞无核到多核;子实层担孢子可含0-4个不等的细胞核,不同时期弹射的担孢子含有的细胞核数量不同。桦纤孔菌担孢子萌发率极低,能萌发的担孢子多为早期弹射的担孢子;培养基也影响担孢子的萌发率,与PDA培养基和CYM培养基相比,桦木屑培养基最适合桦纤孔菌担孢子萌发,萌发率为4.55%。从担孢子萌发的96个菌株中获得了2个不结实菌株和9个结实不产孢菌株,占11.5%,这些菌株间亲和试验出现不同的表现特征,包括形成产孢子实体,产生菌丝纽结,相互融合和相互拮抗等现象,认为桦纤孔菌的有性生殖以次级同宗结合为主,并受交配型基因控制。交配型位点克隆测序后分析发现,桦纤孔菌交配型A位点共14 034 bp,含有一个MIP基因和两组HD1和HD2基因;交配型B位点包含3个疑似信息素受体基因和1个信息素前体编码基因。     

Morphology and evolution of sexual reproduction in the Volvocaceae (Chlorophyta)

Morphological features of sexual reproduction in the Volvocaceae are reviewed, focusing particularly on gametic union and zygote gemination. Both of the two conjugating gametes of the isogamous generaPandorina, Volvulina andYamagishiella bear a tubular mating structure (mating papilla), and plasmogamy is initiated by union of the papillae tips. On zygote germination, a single biflagellate gone cell is released from the zygote wall. Although all the anisogamous and oogamous genera of the Volvocaceae produce “sperm packets” during gametogenesis and a single gone cell at zygote germination, some difference can be recognized in the male gametes. The male gametes ofEudorina bear a tubular cytoplasmic protuberance (putative mating papllla) near the base of the flagella, whereas such a structure recognized at the light microscopic level is not evident inPleodorina andVolvox. Evolution of the sexual reproduction characteristics of volvocacean algae is discussed on the basis of recent cladistic analysis of morphological data as well as of the ribosomal (r) RNA phylogeny and large subunit of the ribulose-1, 5-biphosphate carboxylase/oxygenase(rbcL) gene trees. Dedicated to Emeritus Prof. Hideo Kasaki (Tokyo Metropolitan University) on the occasion of his 77th birthday. Recipient of the Botanical Society Award for Young Scientists, 1994.     

Sexual reproduction in the arum lily family,with emphasis on thermogenicity

Summary Floral thermogenicity, which is found in several representatives of half a dozen angiosperm families, is most pronounced in the Araceae. It is based on the operation of an alternative, cyanide-resistant electron transport chain which, in contrast to the classic cytochrome oxidase system, produces little ATP; most of the energy originally locked up in the respiratory substrate usually starch — is therefore liberated in the form of heat. The biological function of this (biochemically wasteful) system is to release the heat to serve as a volatilizer for the floral odors (often containing aliphatic amines, indole and skatole) that attract the insect pollinators. This makes the survival value of thermogenesis (for the plant species) immediately clear. Thermogenicity is under tight biological control, as demonstrated by the fact that the same ceiling temperature is always reached, regardless of ambient temperature. In Eastern skunk cabbage (Symplocarpus foetidus), which flowers very early in spring, that ceiling is about 20° C, in tropical forms such as Xanthosoma robustum and Philodendron selloum, it lies in the 42°–44° C range. In several instances, e.g., in Arum and in Sauromatum, the voodoo lily, thermogenicity manifests itself as a flare-up of only a few hours' duration, a respiratory explosion that can lead to rates of metabolism that compare favorably with those of a hovering hummingbird. The metabolic peak is always reached at a particular time of day, which is different for the different arum lily species, and thus reduces competition for pollinators. The odors that accompany the heat are also very characteristic, appealing to different pollinator classes and further reducing such competition. In the voodoo lily and in Arum, the primary site for the production of both heat and odor is the naked appendix of the inflorescence, which acts as a specialized osmophore or odor carrier. The first explosion may be followed by another one several hours later, which manifests itself in the floral chamber of the inflorescence and is under strict photoperiodic control. In Sauromatum, the first metabolic explosion is triggered by a plant hormone, originally referred to as calorigen, which originates in the primordia of the staminate flowers and moves from there into the appendix where it exerts its action after a lag-time of about a day — an indication that synthesis of new enzymatic protein (through unblocking of certain genes?) may well be involved. In 1987, calorigen was shown to be identical with salicylic acid. This compound was already known to induce flowering in certain duck-weeds, Lemnaceae, which until recently were regarded as belonging to the same family as arum lilies. In certain water lilies (Nymphaeaceae), thermogenicity is combined with a pollination syndrome very similar to that of Arum and Sauromatum but involving temporary trap flowers.Dedicated to the memory of A.W.H. van Herk, pioneer in the study of arum lilies, superb lecturer, and unselfish human being     

萼花臂尾轮虫有性生殖、种群增长和休眠卵产量间的关系

利用种群指数增长和Logistic增长模型,通过计算机模拟研究了萼花臂尾轮虫有性生殖发生的频率,后代中的混交雌体发率对种群增长和休眠卵产量的影响。在所模拟的参数范围内,随有性生殖发生频率由100％减小到20％,获得了大休眠卵产量所需的混交雌体百分率由9％增大到69％,随密度制约作用的增大（环境容纳量K值由1000减小到100）,该混交雌体百分率由18％增大到69％,休眠卵产量由1072．10降低至133．67,种群的内禀增长率与获得最大休眠卵产量所需的混交雌体百分率间呈曲线相关。当有性生殖发生得越频繁,种群增长所受的密度制约作用较小时,后代中10％－30％的个体为混交雌体时的种群中休眠卵产量较大。     

Coalescent theory for a completely random mating monoecious population

Consider a large random mating monoecious diploid population that has N individuals in each generation. Let us assume that at time 0 a random sample of ninfinity. It is then possible to obtain a generalization of coalescent theory for haploid populations if the distribution of G1 has a finite second moment and E[G(1)(3)]/N-->0 as N-->infinity.     

Costs of reproduction in Nyssa sylvatica: sexual dimorphism in reproductive frequency and nutrient flux

Summary We examined the influence of differential reproductive frequency between the sexes on tertiary (phenotypic) sex ratios in the the dioecious tree Nyssa sylvatica (Nyssaceae). Reproduction was evaluated in relation to sex, size and canopy exposure using flowering data collected from 1229 marked trees over a four year period. For subsets of each population we used data on flower number, fruit crop size, fruit/flower ratios, and individual flower and fruit mass to compare biomass invested in reproductive structures of males and females. We also examined seasonal changes in stem nitrogen and soluble carbohydrate content in relation to flower and fruit production for trees of each sex. Our results indicate that: 1) Male-biased tertiary sex ratios could be explained by more frequent reproduction by male trees; 2) Estimated secondary sex ratios based on sums of all known males and females were not significantly different from 1:1; 3) Flowering frequency of males and females was significantly related to plant size (DBH) and exposure of the canopy to light; 4) Estimtes of reproductive biomass allocation ranged from 1.36 to 10.8 times greater for females relative to males; 5) Flower production was related to stem nutrient status for both sexes, but nutrient depletion and its effect on subsequent flowering was much more pronounced for female trees. We conclude that less frequent flowering by female trees may result from depletion of stored reserves, and that differential flowering frequency in N. sylvatica may ultimately reduce apparent sexual differences in the costs of reproduction.     

Wright-Fisher model of social insects with haploid males and diploid females

An inhomogeneous discrete Markov model is formulated for sexual random mating in finite populations of haploid male and diploid female individuals. This is a Wright-Fisher type of model for social insects. The generations are non-overlapping and of given finite sizes. Bottlenecks are included, allowing different sizes to change from generation to generation. Mutations and selection are included in this exact model for the stochastic process. Computations of the exact Markov model are presented, focussing on the sexually asymmetric genetic drift caused by haplodiploidy.     

松嫩平原碱化草甸星星草种群营养繁殖及有性生殖的数量特征

根据对星星草单优群落独立株丛的大样本取样和对正常花序随机取样的调查与测定,分析了松嫩平原碱化草甸星星草种群的营养繁殖及有性生殖特征．结果表明,星星草种群诸营养繁殖数量性状之间有明显的规律性,丛植株数、丛地上生物量、丛生殖株数及丛生殖株重分别随丛径呈指数形式增加．星星草种群的圆锥花序大多为９—１６个节,花序上各节位上的小穗数量呈γ密度函数分布．随着节位的增高,各节位上的小花数量呈Ｌｏｇｉｓｔｉｃ曲线形式减少．小穗数和小花数均随节位增高呈饱和型指数曲线形式累积增长．     

Isolation of myosin XI genes from the Closterium peracerosum-strigosum-littorale complex and analysis of their expression during sexual reproduction

Myosins comprise a large superfamily of molecular motors that generate mechanical force in ATP-dependent interactions with actin filaments. On the basis of their conserved motor domain sequences, myosins can be divided into at least 17 classes, 3 of which (VIII, XI, XIII) are found in plants. Although full sequences of myosins are available from several species of green plants, little is known about the functions of these proteins. Additionally, sequence information for algal myosin is incomplete, and little attention has been given to the molecular evolution of myosin from green plants. In the present study, the Closterium peracerosum-strigosum-littorale complex was used as a model system for investigating a unicellular basal charophycean alga. This organism has been well studied with respect to sexual reproduction between its two mating types. Three types of partial sequences belonging to class XI myosins were obtained using degenerate primers designed to amplify motor domain sequences. Real-time polymerase chain reaction analysis of the respective myosin genes during various stages of the algal life cycle showed that one of the genes was more highly expressed during sexual reproduction, and that expression was cell-cycle-dependent in vegetatively grown cells.