Soft selection reduces loss of heterozygosity in asexual reproduction

The adaptive value of sexual reproduction is still debated in evolutionary theory. It has been proposed that the advantage of sexual reproduction over asexual reproduction is to promote genetic diversity, to prevent the accumulation of harmful mutations or to preserve heterozygosity. Since these hypothetical advantages depend on the type of asexual reproduction, understanding how selection affects the taxonomic distribution of each type could help us discriminate between existing hypotheses. Here, I argue that soft selection, competition among embryos or offspring in selection arenas prior to the hard selection of the adult phase, reduces loss of heterozygosity in certain types of asexual reproduction. Since loss of heterozygosity leads to the unmasking of recessive deleterious mutations in the progeny of asexual individuals, soft selection facilitates the evolution of these types of asexual reproduction. Using a population genetics model, I calculate how loss of heterozygosity affects fitness for different types of apomixis and automixis, and I show that soft selection significantly reduces loss of heterozygosity, hence increases fitness, in apomixis with suppression of the first meiotic division and in automixis with central fusion, the most common types of asexual reproduction. Therefore, if sexual reproduction evolved to preserve heterozygosity, soft selection should be associated with these types of asexual reproduction. I discuss the evidence for this prediction and how this and other observations on the distribution of different types of asexual reproduction in nature is consistent with the heterozygosity hypothesis.     

Population cycles caused by selection by density dependent competitive interactions

Several animal species have cyclic population dynamics with phase-related cycles in life history traits such as body mass, reproductive rate, and pre-reproductive period. Although many mechanisms have been proposed there is no agreement on the cause of these cycles, and no population equation that deduces both the abundance and the life history cycles from basic ecological constraints has been formulated. Here I deduce a population dynamic equation from the selection pressure of density dependent competitive interactions in order to explain the cyclic dynamics in abundance and life history traits. The model can explain cycles by evolutionary changes in the genotype or by plastic responses in the phenotype. It treats the population dynamic growth rate as an initial condition, and its density independent fundament is Fisher’s (1930, The Genetical Theory of Natural Selection, Oxford: Clarendon) fundamental theorem of natural selection that predicts a hyper-geometrical increase in abundance. The predicted periods coincide with the cyclic dynamics of Lepidoptera, and the Calder hypothesis, which suggests that the period of population cycles is proportional to the 1/4 power of body mass, follows from first principles of the proposed density dependent ecology.     

On selection dynamics for competitive interactions

In this paper, we are interested in an integro-differential model that describe the evolution of a population structured with respect to a continuous trait. Under some assumption, we are able to find an entropy for the system, and show that some steady solutions are globally stable. The stability conditions we find are coherent with those of Adaptive Dynamics.     

The kin selection theory of genomic imprinting and modes of reproduction in the eusocial Hymenoptera

Genomic imprinting is known from flowering plants and mammals but has not been confirmed for the Hymenoptera even though the eusocial Hymenoptera are prime candidates for this peculiar form of gene expression. Here, the kin selection theory of genomic imprinting is reviewed and applied to the eusocial Hymenoptera. The evidence for imprinting in eusocial Hymenoptera with the typical mode of reproduction, involving the sexual production of diploid female offspring, which develop into workers or gynes, and the arrhenotokous parthenogenesis of haploid males, is also reviewed briefly. However, the focus of this review is how atypical modes of reproduction, involving thelytokous parthenogenesis, hybridisation and androgenesis, may also select for imprinting. In particular, naturally occurring hybridisation in several genera of ants may provide useful tests of the role of kin selection in the evolution of imprinting. Hybridisation is expected to disrupt the coadaptation of antagonistically imprinted loci, and thus affect the phenotypes of hybrids. Some of the limited data available on hybrid worker reproduction and on colony sex ratios support predictions about patterns of imprinting derived from kin selection theory.     

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.     

Character compatibility of molecular markers to distinguish asexual and sexual reproduction

Particularly in polyploids, the potential of the high variability of dominant markers such as random amplified polymorphic DNA fragments (RAPDs) and amplified fragment length polymorphisms (AFLPs) in population genetic studies and analysis of breeding systems is reduced due to their dominant nature. In contrast, the criterion of character compatibility is hindered neither by dominance nor by polyploidy as allelic interpretation is not necessary. Character compatibility, which can be used to detect events of genetic exchange (or recombination), is particularly informative if these events are expected to be rare such as in taxa with extensive vegetative reproduction or apomixis. Binary unordered characters such as presence and absence of anonymous DNA markers are incompatible if all four pairwise combinations of character states are present among the individuals studied. Because incompatible character state distributions defy any progenitor–derivative relationship among individuals, they provide strong evidence for genetic exchange. Both the absolute number of incompatible character combinations and the probability of compatibility can be used as a measure of incompatibility. Although these measures may not directly relate to the frequency of genetic exchange, they provide a useful tool to heuristically explore data sets. The most commonly used input for multivariate analyses and analysis of molecular variance in population genetic studies of (dis)similarity of marker distributions are amalgamates of mutation and recombination. Character compatibility can be used to complement these traditional methods of analysis. Advantages and disadvantages of character incompatibility relative to multilocus analysis of modes of reproduction and population genetics are demonstrated with data from RAPDs, isozymes, and restriction fragment length polymorphisms (RFLPs) of the nuclear ribosomal and chloroplast genome.     

Environmental stimulants to asexual reproduction in the planarian,Dugesia dorotocephala

Studies of environmental factors that stimulate asexual reproduction in the planarian,Dugesia dorotocephala (Woodworth), feeding onCulex pipiens quinquefasciatus Say revealed that unchlorinated well and irrigation waters were superior to buffered aqueous media for fission. Constant darkness also was superior to constant light; although, a 12/12-hr photoperiod also allowed good asexual reproduction. Fluctuating dissolved oxygen levels between 3 and 8 ppm apparently stimulated fission. A coarse gravel substrate was superior to smooth plastic. Daily temperature increases of 0.8°C/day between 22.5° and 28.9°C also stimulated fission. Fission rates decreased with increased densities ofD. dorotocephala in unfiltered aquaria, contrary to earlier results obtained in filtered aquaria. The addition of the ostracods,Cypridopsis vidua (Müller) andPhysocypria pustulosa Sharpe, to a diet of mosquito larvae increased fission. Brine shrimp (Artemia sp.) and red tubifex worms (Tubifex sp.) were not as suitable as mosquito larvae for food, apparently because the planaria had difficulty capturing these prey.D. dorotocephala was tolerant of salinities between 250–1,000 ppm, but feeding activity was impaired above and below this range. Mechanical sectioning of individuals was not a practical method for mass production; although if sectioning were coupled with media filtration, some reproductive acceleration might result.

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Résumé Les études sur les facteurs de l'environnement qui stimulent la reproduction asexuée chez la planaire,Dugesia dorotocephala (Woodworth), se nourrissant deCulex pipiens quinquefasciatus Say ont mis en évidence que les puits d'eau ordinaires et les eaux d'irrigation sont supérieurs à d'autres milieux aqueux pour la division; et que le noir total est supérieur à la lumière totale bien qu'une photopériode de 12/12 soit aussi un optimum. Des niveaux variables d'oxygène dissous entre 3 et 8 ppm semblent stimuler la reproduction. Un substrat de gros gravier est supérieur à une surface de plastique lisse et des augmentations quotidiennes de température de 0,8°C/jour entre 22,5° et 28,9°C ont stimulé la partition. La fréquence de multiplication a été diminuée par l'augmentation de la densité enDugesia dans les aquariums non-filtrés, contrairement aux résultats antérieurs dans les aquariums filtrés. L'addition des ostracodes,Cypridopsis vidua (Müller) etPhysocypria pustulosa Sharpe à un régime alimentaire à base de larves de moustiques a augmenté la reproduction. Les crevettes de saumure,Artemia sp., et les vers rougesTubifex sp., ne conviennent pas autant que les larves de moustiques comme aliment, à cause des difficultés, semble-t-il, de capture de ces proies.D. dorotocephala a supporté des salinités entre 250–1,000 ppm, mais l'activité alimentaire a été diminuée au-dessus et audessous de ces extrêmes. Le sectionnement mécanique des individus n'est pas une méthode pratique pour une production massive, bien que, associé à une filtration, il soit possible d'obtenir une accélération de la reproduction.

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Research partially supported by funds from a special appropriation for mosquito research authorized by the California State Legislature during 1975.     

The cost of switching from asexual to sexual reproduction in an aphid

The fecundity of asexual and sexual morphs of Sitobion avenae (F.) is compared. The cost of sex is quantified in terms of the reduction in numbers of offspring produced. The cost is much greater than the halving of fecundity that is usually assumed to be associated with the production of males.

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Résumé Les fécondités des virginipares ailés, des gynopares ailés et des ovipares d'un même clone de Sitobion avenae ont été déterminées à 12°C. Bien que les types qui ayant donné naissance à des sexués aient eu une fécondité initiale plus élevée que celle de la lignée asexuée, celle-ci a été significativement plus féconde après 15 jours de reproduction.Les résultats du laboratoire ont été utilisés pour évaluer la fécondité de lignées sexuées et asexuées pendant quatre générations à 12°C. La reproduction parthénogénétique a présenté un avantage numérique sur la reproductions sexuée dans le rapport 320/1. La reproduction sexuée a été accompagnée d'une diminution significative de la fécondité potentielle. Bien que la reproduction sexuée de S. avenae soit avantageuse dans les régions où les hivers sont durs, parce que les oeufs sont résistants au froid, les lignées asexuées présentent un avantage considérable en ce qui concerne la fécondité potentielle pour l'augmentation de la population en automne et elles conservent de plus la capacité d'un accroissement numérique ultérieur si l'hiver est doux.

     

Social phenotype extended to communities: Expanded multilevel social selection analysis reveals fitness consequences of interspecific interactions

In social species, fitness consequences are associated with both individual and social phenotypes. Social selection analysis has quantified the contribution of conspecific social traits to individual fitness. There has been no attempt, however, to apply a social selection approach to quantify the fitness implications of heterospecific social phenotypes. Here, we propose a novel social selection based approach integrating the role of all social interactions at the community level. We extended multilevel selection analysis by including a term accounting for the group phenotype of heterospecifics. We analyzed nest activity as a model social trait common to two species, the lesser kestrel (Falco naumanni) and jackdaw (Corvus monedula), nesting in either single‐ or mixed‐species colonies. By recording reproductive outcome as a measure of relative fitness, our results reveal an asymmetric system wherein only jackdaw breeding performance was affected by the activity phenotypes of both conspecific and heterospecific neighbors. Our model incorporating heterospecific social phenotypes is applicable to animal communities where interacting species share a common social trait, thus allowing an assessment of the selection pressure imposed by interspecific interactions in nature. Finally, we discuss the potential role of ecological limitations accounting for random or preferential assortments among interspecific social phenotypes, and the implications of such processes to community evolution.     

用珍珠洋葱探究植物生理知识的组合实验

以珍珠洋葱为实验材料,以组合实验的形式探究无性繁殖与植物生长的知识,以培养学生解决问题的综合能力。     

Convergence to constant equilibrium for a density-dependent selection model with diffusion

We consider the classical single locus two alleles selection model with diffusion where the fitnesses of the genotypes are density dependent. Using a theorem of Peter Brown, we show that in a bounded domain with homogeneous Neumann boundary conditions, the allele frequency and population density converge to a constant equilibrium lying on the zero population mean fitness curve. The results agree with the case without diffusion obtained by Selgrade and Namkoong. Frequency and density dependent selection is also considered.Research partially supported by NSF grant DMS-8601585     

Ant queens adjust egg fertilization to benefit from both sexual and asexual reproduction

An enduring problem in evolutionary biology is the near ubiquity of sexual reproduction despite the inherent cost of transmitting only half the parent's genes to progeny. Queens of some ant species circumvent this cost by using selectively both sexual reproduction and parthenogenesis: workers arise from fertilized eggs, while new queens are produced by parthenogenesis. We show that queens of the ant Cataglyphis cursor maximize the transmission rate of their genes by regulating the proportion of fertilized and parthenogenetic eggs laid over time. Parthenogenetic offspring are produced in early spring, when workers raise the brood into sexuals. After the mating period, queens lay mostly fertilized eggs that will be reared as the non-reproductive caste.     

Using multilevel models to quantify heterogeneity in resource selection

Models of resource selection are being used increasingly to predict or model the effects of management actions rather than simply quantifying habitat selection. Multilevel, or hierarchical, models are an increasingly popular method to analyze animal resource selection because they impose a relatively weak stochastic constraint to model heterogeneity in habitat use and also account for unequal sample sizes among individuals. However, few studies have used multilevel models to model coefficients as a function of predictors that may influence habitat use at different scales or quantify differences in resource selection among groups. We used an example with white-tailed deer (Odocoileus virginianus) to illustrate how to model resource use as a function of distance to road that varies among deer by road density at the home range scale. We found that deer avoidance of roads decreased as road density increased. Also, we used multilevel models with sika deer (Cervus nippon) and white-tailed deer to examine whether resource selection differed between species. We failed to detect differences in resource use between these two species and showed how information-theoretic and graphical measures can be used to assess how resource use may have differed. Multilevel models can improve our understanding of how resource selection varies among individuals and provides an objective, quantifiable approach to assess differences or changes in resource selection. © 2011 The Wildlife Society.     

Site-directed selection of oligonucleotide antagonists by competitive elution

Oligonucleotide ligands that bind a protein or a small molecule of interest are readily isolated by in vitro selection and amplification of rare sequences from combinatorial libraries of sequence-randomized oligonucleotides (Gold et al., 1995). Classic systematic evolution of ligands by exponential enrichment (SELEX) protocols are affinity based (Tuerk and Gold, 1990), but because many problems and applications require antagonists, protocols for selecting inhibitors are both desirable and valuable. A widely applicable approach for isolating inhibitors is competitive elution with a molecule that binds the targeted molecule's active or binding site. We have used this approach to isolate antagonists of wheat germ agglutinin (WGA) from a library of 2'NH2-pyrimidine, 2'OH-purine oligonucleotides by elution with N N' N"-triacetylchitotriose, (GlcNAc)3. The highest affinity aptamers have equilibrium dissociation constants of 1 nM-20 nM for WGA, a 10(3)-10(4)-fold improvement relative to (GlcNAc)3, and unlike the carbohydrate, are highly specific. In addition to competing for binding with (GlcNAc)3, aptamers inhibit WGA-mediated agglutination of sheep erythrocytes, demonstrating that they are able to compete with natural ligands presented on the surfaces of cells. These results illustrate the feasibility of isolating high-affinity, high-specificity antagonists by competitive elution with low molecular weight, relatively low-affinity, and low-specificity small molecules.     

Locomotion,asexual reproduction,and killing of corals by the corallimorpharian Corynactis californica

The rates at which some sedentary coelenterates move across substrates, spread by asexual reproduction, and competitively damage their neighbors may greatly affect their ability to survive on hard marine substrates. However, almost nothing is known about these processes in the Corallimorpharia. In this study, locomotory rates varied widely between members of a single clone of the corallimorpharian Corynactis californica. Polyp movement rates (0–14 mm month^–1) were slower than those of many Actiniaria, possibly due to the lack of basilar muscles in corallimorpharians. Rates of asexual reproduction by C. californica polyps in the laboratory varied 10-fold among 8 clones. The minimal time to double the number of polyps per clone (2 months) was intermediate to doubling times known for Actiniaria and Scleractinia. The rate at which polyps killed individuals of the scleractinian Astrangia lajollaensis also varied widely within and between clones. Asexual reproductive rate correlated positively with coral killing rate.     

Verticillium dahliae Asp1 regulates the transition from vegetative growth to asexual reproduction by modulating microtubule dynamic organization

Verticillium dahliae is a devastating pathogenic fungus that causes severe vascular wilts in more than 400 dicotyledonous plants. The conidiation of V. dahliae in plant vascular tissues is the key strategy for its adaptation to the nutrient-poor environment and is required for its pathogenicity. However, it remains unclear about the regulatory mechanism of conidium production of V. dahliae in vascular tissues. Here, we found that VdAsp1, encoding an inositol polyphosphate kinase, is indispensable for the pathogenicity of V. dahliae. Loss of VdAsp1 function does not affect the invasion of the host, but it impairs the colonization and proliferation in vascular tissues. The ΔVdAsp1 mutant shows defective initiation of conidiophore formation and reduced expression of genes associated with the central developmental pathway. By live-cell imaging, we observed that some of ΔVdAsp1 mutant hyphae are swollen, and microtubule arrangements at the apical region of these hyphae are disorganized. These results indicate that VdAsp1 regulates the transition from vegetative growth to asexual reproduction by modulating microtubule dynamic organization, which is essential for V. dahliae to colonize and proliferate in vascular tissues. These findings provided a potential new direction in the control of vascular wilt pathogen by targeting conidium production in vascular tissues.     

Lectin-carbohydrate interactions studied by a competitive enzyme inhibition assay

The principle of competitive binding assay in combination with an immobilized lectin (concanavalin A), in close proximity to an oxygen sensor, has been used to quantify carbohydrates and to determine association constants for lectin-carbohydrate interactions. Methyl α-d-mannopyranoside was determined down to 0.5 μg/ml. K_a (maltose) and K_a (maltotriose) was found to be 2.1 × 10³ and 1.7 × 10³m^?1, respectively, which are comparable to values quoted in the literature of approximately 2.8 × 10³m^?1 for both maltose and maltotriose. Furthermore, the estimation of the bonus effect, due to multipoint attachment, for a low-molecular-weight dextran is discussed.     

From patterns to processes and back: analysing density-dependent responses to an abiotic stressor by statistical and mechanistic modelling

Our knowledge about population-level effects of abiotic stressors is limited, largely due to lack of appropriate time-series data. To analyse interactions between an abiotic stressor and density-dependent processes, we used experimental time-series data for stage-structured populations (the blowfly Lucilia sericata) exposed to the toxicant cadmium through 20 generations. Resource limitation results in competition both in the larval and the adult stages. The toxicant has only negative effects at the organism level, but nevertheless, there were positive population-level effects. These are necessarily indirect, and indicate overcompensatory density-dependent responses. A non-parametric model (generalized additive model) was used to investigate the density-dependent structures of the demographic rates, without making assumptions about the functional forms. The estimated structures were used to develop a parametric model, with which we analysed effects of the toxicant on density-dependent and density-independent components of the stage-specific demographic rates. The parameter estimates identified both synergistic and antagonistic density-toxicant interactions. It is noteworthy that the synergistic interaction occurred together with a net positive effect of the toxicant. Hence, the effects of such interactions should be considered together with the capacity for compensatory responses. The combination of the two modelling approaches provided new insight into mechanisms for compensatory responses to abiotic stressors.     

The response to fire by two eusocial bee species

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