A mutation in the cAMP signaling pathway affects sexual development of Dictyostelium discoideum

Amoebae of cellular slime molds have two developmental modes, asexual fruiting body formation and sexual macrocyst formation. How developmental choice is made is an interesting subject of wide importance. Light exposure and dry conditions are favorable for asexual development, while conditions of darkness and high humidity are so for sexual development. In Dictyostelium discoideum , the latter conditions enhance zygote formation, which determines the fate of surrounding cells for sexual development. Here, a mutant (TMC1) defective in the post-fusion aggregation of cells during sexual development is described. This mutant is also aggregationless in asexual development, and the level of cyclic adenosine monophosphate (cAMP) receptor is reduced. Correspondingly, a series of existing mutants with defects in cAMP signaling pathways showed the same sexual phenotype as TMC1. These results suggest that molecular mechanisms of development are shared by the two alternative developmental modes.     

The Tangled Bank: The maintenance of sexual reproduction through competitive interactions

The maintenance of sexual reproduction is discussed using a model based on the familiar Lotka-Volterra competition equations. Both the equilibrium and the stability conditions that allow a sexual population to resist invasion by a single asexual clone are considered. The equilibrium conditions give results similar to previous models: When the cost of sex, within phenotype niche width, and environmental variance are low, the sexual population coexists with the asexual clone and remains at a high density. However, the asexual clone is never completely excluded. Analysis of the stability conditions shows a different picture: The introduction of an asexual clone considerably reduces the stability of the community. However, owing to its larger total niche width, the sexual population exists partly in a “competitor-free space” where the asexual clone has almost no influence on the outcome of the interactions. Therefore the asexual clone is less stable than the sexual population and has a higher probability of extinction. In contrast, the sexual population does not become extinct, since the extreme phenotypes remain at a stable, though low, density, and the central phenotypes, where stability is low, are recreated every generation through recombination. I therefore conclude that the ecological conditions under which sexual reproduction is favored over asexual reproduction are fairly easily attained and are more general than previous analyses had suggested.     

On the fate of sexual traits under asexuality

Environmental shifts and life‐history changes may result in formerly adaptive traits becoming non‐functional or maladaptive. In the absence of pleiotropy and other constraints, such traits may decay as a consequence of neutral mutation accumulation or selective processes, highlighting the importance of natural selection for adaptations. A suite of traits are expected to lose their adaptive function in asexual organisms derived from sexual ancestors, and the many independent transitions to asexuality allow for comparative studies of parallel trait maintenance versus decay. In addition, because certain traits, notably male‐specific traits, are usually not exposed to selection under asexuality, their decay would have to occur as a consequence of drift. Selective processes could drive the decay of traits associated with costs, which may be the case for the majority of sexual traits expressed in females. We review the fate of male and female sexual traits in 93 animal lineages characterized by asexual reproduction, covering a broad taxon range including molluscs, arachnids, diplopods, crustaceans and eleven different hexapod orders. Many asexual lineages are still able occasionally to produce males. These asexually produced males are often largely or even fully functional, revealing that major developmental pathways can remain quiescent and functional over extended time periods. By contrast, for asexual females, there is a parallel and rapid decay of sexual traits, especially of traits related to mate attraction and location, as expected given the considerable costs often associated with the expression of these traits. The level of decay of female sexual traits, in addition to asexual females being unable to fertilize their eggs, would severely impede reversals to sexual reproduction, even in recently derived asexual lineages. More generally, the parallel maintenance versus decay of different trait types across diverse asexual lineages suggests that neutral traits display little or no decay even after extended periods under relaxed selection, while extensive decay for selected traits occurs extremely quickly. These patterns also highlight that adaptations can fix rapidly in natural populations of asexual organisms, in spite of their mode of reproduction.     

Existence of two sexual races in the planarian species switching between asexual and sexual reproduction

In certain planarian species that are able to switch between asexual and sexual reproduction, determining whether a sexual has the ability to switch to the asexual state is problematic, which renders the definition of sexuals controversial. We experimentally show the existence of two sexual races, acquired and innate, in the planarian Dugesia ryukyuensis. Acquired sexuals used in this study were experimentally switched from asexuals. Inbreeding of acquired sexuals produced both innate sexuals and asexuals, but inbreeding of innate sexuals produced innate sexuals only and no asexuals. Acquired sexuals, but not innate sexuals, were forced to become asexuals by ablation and regeneration (asexual induction). This suggests that acquired sexuals somehow retain asexual potential, while innate sexuals do not. We also found that acquired sexuals have the potential to develop hyperplastic and supernumerary ovaries, while innate sexuals do not. In this regard, acquired sexuals were more prolific than innate sexuals. The differences between acquired and innate sexuals will provide a structure for examining the mechanism underlying asexual and sexual reproduction in planarians.     

圆盘菌科无性型有性生殖能力评估

本文评估了分离自子囊孢子和分生孢子的圆盘菌科无性型有性生殖能力。结果表明,由子囊孢子和分生孢子萌发获得的菌株的有性生殖能力具有显著差异,这可能决定于不同的遗传特性。该试验支持了一种假说：仅能进行无性生殖的无性型菌株,很可能来源于逐渐丧失了有性生殖能力的全型菌株。     

Antagonism between sexual and natural selection in experimental populations of Saccharomyces cerevisiae

Trade-offs between life-history components are a central concept of evolution and ecology. Sexual and natural selection seem particularly apt to impose antagonistic selective pressures. When sex is not integrated into reproduction, as in Saccharomyces cerevisiae, natural selection can impair or even eliminate it. In this study, a genetic trade-off between the sexual and asexual phases of the yeast life cycle was suggested by sharp declines in the mating and sporulation abilities of unrelated genotypes that were propagated asexually in minimal growth medium and in mice. When sexual selection was applied to populations that had previously evolved asexually, sexual fitness increased but asexual fitness declined. No such negative correlation was observed when sexual selection was applied to an ancestral strain: sexual and asexual fitness both increased. Thus, evolutionary history affected the evolution of genetic correlations, as fitness increases in a population already well adapted to the environment were more likely to come at the expense of sexual functions.     

Extreme tolerance to environmental stress of sexual and parthenogenetic resting eggs of Eucypris virens (Crustacea,Ostracoda)

1. The freshwater ostracod (Ostracoda), Eucypris virens, is commonly found in European temporary pools, where its long‐term persistence completely relies on the build‐up of resting egg banks. Extreme tolerance of dormant eggs and seeds is widely assumed, but freshwater ostracod eggs are relatively poorly studied. The study of ostracod resting eggs is of particular relevance as it may yield the key to understanding the distribution of the sexes in many species capable of both sexual and asexual reproduction. 2. We assessed the tolerance of dried resting eggs produced by females originating from three populations with males and three all‐female E. virens populations. Hatching time and success was compared between control eggs and eggs exposed to one of seven ecologically relevant stressors: digestive enzymes, high salinity, deep freezing, hydration, UV‐B radiation, hypoxia and insecticide treatment. 3. None of the stressors reduced significantly the viability of either sexual or asexual eggs. When compared with the reproductive mode–specific controls, exposure to UV‐B radiation had a mild impact on the survival of sexual and asexual eggs (?16.8 and ?22.4%, respectively), but this was only significant for asexual eggs. These results point to an extreme tolerance of E. virens resting eggs and have important implications for the ecology and evolution of the species. 4. The timing of hatching was not affected by the stress treatment, except for UV‐B radiation. A marginally significant delay in hatching response was observed for UV‐B‐radiated eggs when compared to the overall mean, but this treatment effect was absent when compared with the reproductive mode–specific controls. 5. The populations with males produced eggs that hatched on average earlier (?1.5 days at 17 °C) and were more successful (+26%) than asexual eggs. Due to the limited number of populations and the population‐specific origin and age of the eggs, the possibility due to the differences in age and origin of the resting eggs, or to variations in local conditions, cannot be ruled out.     

昆虫抑性欲素研究进展

抑性欲素(anti-aphrodisiac)是雄性昆虫在交配过程中留给雌性的气味物质,该物质可以降低交配过的雌性对其它同种雄性的吸引力。本文综述了有抑性欲素研究报道的昆虫种类、抑性欲素对蝶类性行为的影响以及赤眼蜂对昆虫抑性欲素利用情况的研究进展。     

Temporal differentiation and spatial coexistence of sexual and facultative asexual lineages of an aphid species at mating sites

Cases of coexisting sexual and asexual relatives are puzzling, as evolutionary theory predicts that competition for the same ecological niches should lead to the exclusion of one or the other population. In the cyclically parthenogenetic aphid, Rhopalosiphum padi, sexual and facultative asexual lineages are admixed in space at the time of sexual reproduction. We investigated how the interaction of reproductive mode and environment can lead to temporal niche differentiation. We demonstrated theoretically that differential sensitivity of sexual and facultatively asexual aphids to an environmental parameter (mating host suitability) shapes the two strategies: whereas the sexual lineages switch earlier to the production of sexual forms, the facultative asexual lineages delay and spread out their investment in sexual reproduction. This predicted pattern of niche specialization is in agreement with the temporal structure revealed in natura by demographic and genetic data. We propose that partial loss of sex by one pool of aphids and subsequent reduction in gene flow between lineages may favour temporal specialization through disruptive selection.     

The conservation of redundancy in genetic systems: effects of sexual and asexual reproduction

The relationship between probability of survival and the number of deleterious mutations in the genome is investigated using three different models of highly redundant systems that interact with a threatening environment. Model one is a system that counters a potentially lethal infection; it has multiple identical components that act in sequence and in parallel. Model two has many different overlapping components that provide three-fold coverage of a large number of vital functions. The third model is based on statistical decision theory: an ideal detector, following an optimum decision strategy, makes crucial decisions in an uncertain world. The probability of a fatal error is reduced by a redundant sampling system, but the chance of error rises as the system is impaired by deleterious mutations. In all three cases the survival profile shows a synergistic pattern in that the probability of survival falls slowly and then more rapidly. This is different than the multiplicative or independent survival profile that is often used in mathematical models. It is suggested that a synergistic profile is a property of redundant systems. Model one is then used to study the conservation of redundancy during sexual and asexual reproduction. A unicellular haploid organism reproducing asexually retains redundancy when the mutation rate is very low (0001 per cell division), but tends to lose high levels of redundancy if the mutation rate is increased (001 to 01 per cell division). If a similar unicellular haploid organism has a sexual phase then redundancy is retained for mutation rates between 0001 and 01 per cell division. The sexual organism outgrows the asexual organism when the above mutation rates apply. If they compete for finite resources the asexual organism will be extinguished. Variants of the sexual organism with increased redundancy will outgrow those with lower levels of redundancy and the sexual process facilitates the evolution of more complex forms. There is a limit to the extent that complexity can be increased by increasing the size of the genome and in asexual organisms this leads to progressive accumulation of mutations with loss of redundancy and eventual extinction. If complexity is increased by using genes in new combinations, the asexual form can reach a stable equilibrium, although it is associated with some loss of redundancy. The sexual form, by comparison, can survive, with retention of redundancy, even if the mutation rate is above one per generation. The conservation and evolution of redundancy, which is essential for complexity, depends on the sexual process of reproduction.     

Phenotypic variation,sexual reproduction and evolutionary population dynamics

I studied the effects of introducing phenotypic variation into a well-known single species model for a population with discrete, non-overlapping generations. The phenotypes differed in their dynamic behaviour. The analysis was made under the assumption that the population was in an evolutionary stable state. Differences in the timing of the competitive impacts of the phenotypes on each other had a strong simplifying effect on the dynamics. This result could also be applied to competition between species. The effect of sexual reproduction on the dynamics of the population was analysed by assuming the simplest genetic model of one locus with two alleles. Sexual reproduction made the system much more stable in the (mathematical) sense that the number of attractors was reduced and their basins of attraction enlarged. In a dominant system sex tended to increase the frequency of the recessive allele, and in an overdominant system it induced gene frequencies of 1/2. Whether the attractors in the dominant system tended to be simpler or more complex than the attractors in the asexual system depended on the phenotype of the recessive homozygote. The overdominant sexual system tended to have simpler dynamics than the corresponding asexual population. A 2-locus model was used to study whether sexuals can invade an asexual population and vice versa. One locus coded for sexual and asexual reproduction, while the other coded for the dynamics. Enhanced stability through sexual reproduction seemed to be the reason why there was a clear asymmetry favouring sex in this evolutionary context.     

Novel distribution pattern between coexisting sexual and obligate asexual variants of the true estuarine macroalga Ulva prolifera

Where sexual and asexual forms coexist within a species, the asexuals are often found to be prevalent in marginal habitats. This asexual distribution pattern has received evolutionary attention linked to the paradox of sex. In many marine species, there is a distributional trend of asexual modes being more common in lower salinity waters regarded as the ecogeographic marginal, being explained by negative effects of low salinities on sexual reproductive success. However, the distribution pattern of estuarine species recently adapted to low salinity waters has remained unknown. The brackish macroalga Ulva prolifera being a major benthic component of estuarine ecosystems includes a sexual variant and obligate asexual variants by means of motile spores. We examined the sexual–asexual distribution pattern of this alga along a salinity gradient in river estuaries. Surprisingly, opposite to the distributional trend of marine organisms, the results clearly showed the persistent predominance of sexuals in the lower salinity reaches than the asexuals. In addition, a frequent alternating of dioecious gametophytes and sporophytes in the sexual population was observed, suggesting the sexual reproductive process would be robustly performed in the low salinity waters. Considering U. prolifera had evolved from the ancestral marine species to become a true estuarine species of which the core habitat is the low salinity reaches, in a broad sense its sexual–asexual distribution pattern would be involved in asexual marginal occupations of the species range previously reported in other organisms. Based on the frozen niche variation model, we can give a concise explanation for the evolutionary process of this pattern; multiple asexuals with frozen genotypic variation had arisen from sexual ancestors undergoing low salinity adaptation and share the estuarine habitat with the sexuals at present.     

Sexual induction in Volvox carteri f. nagariensis by aldehydes

Summary Sexual induction in the Gone 12 mutant of Volvox carteri can be achieved by shortterm treatment with glutardialdehyde or formaldehyde, followed by capture of the aldehyde by means of amino acids at slightly acidic pH. The same effect is obtained by exposure to anthranilic acid formalide, the condensation product of 2-amino benzoic acid with formaldehyde, at low concentration for several minutes. This is in contrast to the prolonged exposure required by the specific glycoprotein inducer. In both situations the asexual reproductive cells are affected in such a way that they change their pattern of cleavage to form sexual embryos rather than asexual ones. Thus, besides the natural messenger molecule, a physical (UV light, heat) or molecular shock may trigger the chain reaction leading to expression of sexual induction.     

Obligate asex in a rotifer and the role of sexual signals

Transitions to asexuality have occurred in many animals and plants, yet the biological mechanisms causing such transitions have often remained unclear. Cyclical parthenogens, such as cladocerans, rotifers or aphids often give rise to obligate asexual lineages. In many rotifers, chemical signals that accumulate during population crowding trigger the induction of sexual stages. In this study, I tested two hypotheses on the origin of obligate parthenogenesis in the rotifer Brachionus calyciflorus: (i) that obligate parthenogens have lost the responsiveness to the sexual signal; and (ii) that obligate parthenogens have lost the ability to produce the sexual signal. Pairwise cross-induction assays among three obligate parthenogenetic strains and two cyclically parthenogenetic (sexual) strains were used to test these hypotheses. I found that obligate parthenogens can induce sexual reproduction in sexual strains, but not vice versa. This demonstrates that obligate parthenogens do still produce the sexual signal, but have lost responsiveness to that signal.     

Direct and indirect selection in moth pheromone evolution: population genetical simulations of asymmetric sexual interactions

Female moths generally use pheromones to attract males. Normally, all females in a population produce a specific chemical blend with only a limited variance, and the local males are highly attracted to this blend. To better understand the direct and indirect selective forces acting on this communication system, where, unusually, it is the reproductively limited sex that signals for matings, a population genetical model has been constructed and numerically analysed. Basic to the model is the assumption that the pheromone attraction system functions asymmetrically, leading to strong sexual selection between males but no direct sexual selection between females. Evolutionary simulations using the model show that sexual selection in males causes an indirect stabilizing selection on the pheromone blends produced by females. Thus, a more narrow range of pheromone variation is selected for, even in the absence of female sexual selection. The strength of the selection is analysed, and it is suggested that this indirect stabilizing selection becomes particularly important in situations where geographically adjacent populations have evolved different pheromone blends.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 117–123.     

有性生殖对栗疫病菌群体结构的影响

采用RAPD方法对来源于栗疫病菌8个不同子囊壳的子囊孢子后代和无性生殖的对照群体各23个菌株进行了群体结构的比较。从RAPD随机引物中筛选出扩增多态性丰富的4条引物,共扩增出条带73条,多态性检测率为100%。研究结果表明,在8个子囊壳和无性生殖群体中的基因多样性,64.27%由群体内部引起,只有35.73%的多样性由群体之间的基因差异引起。各子囊壳群体间存在的基因流动很小(Nm=0.8994)。有性群体和无性群体之间的遗传距离为0.1389,基因流动值为3.4212,说明子囊壳群体和无性生殖群体之间存在一定的系统关系。分析表明栗疫病菌子囊孢子后代在自然界的传播对自然界的病菌的多样性起重要的作用。     

Parthenogenetic female populations in the brown alga Scytosiphon lomentaria (Scytosiphonaceae,Ectocarpales): decay of a sexual trait and acquisition of asexual traits

In isogamous brown algae, the sexuality of populations needs to be tested by laboratory crossing experiments, as the sexes of gametophytes are morphologically indistinguishable. In some cases, gamete fusion is not observed and the precise reproductive mode of the populations is unknown. In the isogamous brown alga Scytosiphon lomentaria in Japan, both asexual (gamete fusion is unobservable) and sexual populations (gamete fusion is observable) have been reported. In order to elucidate the reproductive mode of asexual populations in this species, we used PCR‐based sex markers to investigate the sex ratio of three asexual and two sexual field populations. The markers indicated that the asexual populations consisted only of female individuals, whereas sexual populations are composed of both males and females. In culture, female gametes of most strains from asexual populations were able to fuse with male gametes; however, they had little to no detectable sexual pheromones, significantly larger cell sizes, and more rapid parthenogenetic development compared to female/male gametes from sexual populations. Investigations of sporophytic stages in the field indicated that alternation of gametophytic and parthenosporophytic stages occur in an asexual population. These results indicate that the S. lomentaria asexual populations are female populations that lack sexual reproduction and reproduce parthenogenetically. It is likely that females in the asexual populations have reduced a sexual trait (pheromone production) and have acquired asexual traits (larger gamete sizes and rapid parthenogenetic development).     

Sexual reproduction and diversity: Connection between sexual selection and biological communities via population dynamics

Sexual reproduction is a mysterious phenomenon. Most animals and plants invest in sexual reproduction, even though it is more costly than asexual reproduction. Theoretical studies suggest that occasional or conditional use of sexual reproduction, involving facultative switching between sexual and asexual reproduction, is the optimal reproductive strategy. However, obligate sexual reproduction is common in nature. Recent studies suggest that the evolution of facultative sexual reproduction is prevented by males that coerce females into sexual fertilization; thus, sexual reproduction has the potential to enforce costs on a given species. Here, the effect of sex on biodiversity is explored by evaluating the reproductive costs arising from sex. Sex provides atypical selection pressure that favors traits that increase fertilization success, even at the expense of population growth rates, that is, sexual selection. The strength of sexual selection depends on the density of a given species. Sexual selection often causes strong negative effects on the population growth rates of species that occur at high density. Conversely, a species that reduces its density is released from this negative effect, and so increases its growth rate. Thus, this negative density-dependent effect on population growth that arises from sexual selection could be used to rescue endangered species from extinction, prevent the overgrowth of common species and promote the coexistence of competitive species. Recent publications on sexual reproduction provide several predictions related to the evolution of reproductive strategies, which is an important step toward integrating evolutionary dynamics, demographic dynamics and community dynamics.     

A sex pheromone, protoplast release-inducing protein (PR-IP) inducer, induces sexual cell division and production of PR-IP in Closterium

The sex pheromone protoplast release-inducing protein (PR-IP) inducer and a sexual cell division-inducing pheromone-minus (SCD-IP-minus) that mediates the sexual reproduction of the heterothallic Closterium peracerosum-strigosum-littorale (C. psl) complex were investigated in this study. Recombinant PR-IP inducer produced by yeast cells was prepared and assayed for production of PR-IP and induction of SCD. Both biological activities were observed after treating mating-type plus (mt+) cells with the recombinant pheromone. SCD was induced by exposure to a lower concentration of the same pheromone and by a shorter treatment period with the pheromone than was production of PR-IP. This indicates that the previously characterized PR-IP inducer has both PR-IP-inducing and SCD-inducing activities with mt+ cells, although the inducing mechanisms of the two pheromones differ.