Fatal or Harmless: Extreme Bistability Induced by Sterilizing,Sexually Transmitted Pathogens

Models of sexually transmitted infections have become a fixture of mathematical epidemiology. A common attribute of all these models is treating reproduction and mating, and hence pathogen transmission, as uncoupled events. This is fine for humans, for example, where only a tiny fraction of sexual intercourses ends up with having a baby. But it can be a deficiency for animals in which mating and giving birth are tightly coupled, and mating thus mediates both reproduction and pathogen transmission. Here, we model dynamics of sterilizing, sexually transmitted infections in such animals, assuming structural consistency between the processes of reproduction and pathogen transmission. We show that highly sterilizing, sexually transmitted pathogens trigger bistability in the host population. In particular, the host population can end up in two extreme alternative states, disease-free persistence and pathogen-driven extinction, depending on its initial state. Given that sterilizing, sexually transmitted infections that affect animals are abundant, our results might implicate an effective pest control tactic that consists of releasing the corresponding pathogens, possibly after genetically enhancing their sterilization power.     

Genetic Mapping and Non-Mendelian Segregation of Mating Type Loci in the Oomycete, Phytophthora Infestans

DNA markers linked to the determinants of mating type in the oomycete, Phytophthora infestans, were identified and used to address the genetic basis of heterothallism in this normally diploid fungus. Thirteen loci linked to the A1 and A2 mating types were initially identified by bulked segregant analysis using random amplified polymorphic DNA markers (RAPDs) and subsequently scored in three crosses as RAPD markers, restriction fragment length polymorphisms (RFLPs), single-strand conformational polymorphisms (SSCP), cleaved amplified polymorphisms (CAPS), or allele-specific polymerase chain reaction markers (AS-PCR). All DNA markers mapped to a single region, consistent with a single locus determining both mating types. Long-range restriction mapping also demonstrated the linkage of the markers to one region and delimited the mating type locus to a 100-kb region. The interval containing the mating type locus displayed non-Mendelian segregation as only two of the four expected genotypes were detected in progeny. This is consistent with a system of balanced lethal loci near the mating type locus. A model for mating type determination is presented in which the balanced lethals exclude from progeny those with potentially conflicting combinations of mating type alleles, such as those simultaneously expressing A1 and A2 functions.     

Joint Frequencies of Alleles Determined by Separate Formulations for the Mating and Mutation Systems

A method to calculate joint gene frequencies, which are the probabilities that two neutral genes taken at random from a population have certain allelic states, is developed taking into account the effects of the mating system and the mutation scheme. We assume that the mutation rates are constant in the population and that the mating system does not depend on allelic states. Under either--the condition that mutation rates are symmetric or that the mating unit is large and the mutation rate is small--the general formula is represented by two terms, one for the mating system and the other for the mutation scheme. The term for the mating system is expressed using the coancestry coefficient in the infinite allele model, and the term for the mutation scheme is a function of the eigenvalues and the eigenvectors of the mutation matrix. Several examples are presented as applications of the method, including homozygosity in a stepping-stone model with a symmetric mutation scheme.     

SEXUAL SELECTION,SPACE, AND SPECIATION

A Fisherian model of sexual selection is combined with a diffusion model of mate dispersal to investigate the evolution of assortative mating in a sympatric population. Females mate with one of two types of polygynous males according to a male's display of one of two sex-limited, autosomal traits; these male traits may be associated with differential phenotypic mortalities. Through a Fisherian runaway process, female preferences and male traits can become associated in linkage disequilibrium, leading to patterns of assortative mating. Dispersing males, whose rate of movement is dependent on mating success, carry female preference genes with them, and displaced males thereby produce daughters with preference genes for their respective traits in locally higher than average frequencies. The reduced diffusion of the more preferred males permits the success of other male types in adjacent areas. Thus, mating-success dependent diffusion, when coupled with the rapid divergence in phenotypes possible under the Fisher process, can lead to the coexistence of two female preferences and two male traits in sympatry. We argue that many existing approaches to sympatric speciation fail to explain observed male polymorphisms because they exclude explicit spatial structure from their speciation models.     

A single locus model of selection in permanent translocation heterozygotes

A model for evolution at a single locus in permanent translocation heterozygotes is described. It is also applicable to other permanent structural heterozygotes that possess the mating systems discussed. Recombination occurs between the locus and the chromosomes, which are of two types. The mating system includes selfing and random mating. When recombination is rare, selection will result in almost complete fixation on the single most fit genotype present in a population, regardless of the frequency of selfing. This provides a possible explanation for the ecological success of permanent translocation heterozygotes in some groups of organisms, like Oenothera. It is not necessary to postulate that their success is the result of hybrid vigor. Furthermore, a complete lack of recombination is not necessary for explaining the observed association of alleles with particular segmental arrangements. A small amount of recombination is consistent with the observation that different segmental arrangements often carry different alleles.     

Central roles of small GTPases in the development of cell polarity in yeast and beyond.

Summary: The establishment of cell polarity is critical for the development of many organisms and for the function of many cell types. A large number of studies of diverse organisms from yeast to humans indicate that the conserved, small-molecular-weight GTPases function as key signaling proteins involved in cell polarization. The budding yeast Saccharomyces cerevisiae is a particularly attractive model because it displays pronounced cell polarity in response to intracellular and extracellular cues. Cells of S. cerevisiae undergo polarized growth during various phases of their life cycle, such as during vegetative growth, mating between haploid cells of opposite mating types, and filamentous growth upon deprivation of nutrition such as nitrogen. Substantial progress has been made in deciphering the molecular basis of cell polarity in budding yeast. In particular, it becomes increasingly clear how small GTPases regulate polarized cytoskeletal organization, cell wall assembly, and exocytosis at the molecular level and how these GTPases are regulated. In this review, we discuss the key signaling pathways that regulate cell polarization during the mitotic cell cycle and during mating.     

Aspergillus genomes: secret sex and the secrets of sex

The genomic sequences of three species of Aspergillus, including the model organism A. nidulans (which is homothallic: having no differentiated mating types, a strain being able to cross with itself), suggest that A. fumigatus and A. oryzae, considered to be asexual, might in fact be heterothallic (having two differentiated mating types, a strain being able to cross only with strains of opposite mating type). The genomic data have implications for the understanding of the evolution and the mechanism of sexual reproduction in this genus. We propose a model of epigenetic heterothallism to account for the reproductive patterns observed in Aspergillus nidulans.     

On the maintenance of bird song dialects

Many bird species, especially song birds but also for instance some hummingbirds and parrots, have noted dialects. By this we mean that locally a particular song is sung by the majority of the birds, but that neighbouring patches may feature different song types. Behavioural ecologists have been interested in how such dialects come about and how they are maintained for over 45 years. As a result, a great deal is known about different mechanisms at play, such as dispersal, assortative mating and learning of songs, and there are several competing hypotheses to explain the dialect patterns known in nature. There is, however, surprisingly little theoretical work testing these different hypotheses at present. We analyse the simplest kind of model that takes into account the most important biological mechanisms, and in which one may speak of dialects: a model in which there are but two patches, and two song types. It teaches us that a combination of little dispersal and strong assortative mating ensures dialects are maintained. Assuming a simple, linear frequency-dependent learning rule has little effect on the maintenance of dialects. A nonlinear learning rule, however, has dramatic consequences and greatly facilitates dialect maintenance. Adding fitness benefits for singing particular songs in a given patch also has a great impact. Now rare song types may invade and remain in the population.     

Central Roles of Small GTPases in the Development of Cell Polarity in Yeast and Beyond

Summary: The establishment of cell polarity is critical for the development of many organisms and for the function of many cell types. A large number of studies of diverse organisms from yeast to humans indicate that the conserved, small-molecular-weight GTPases function as key signaling proteins involved in cell polarization. The budding yeast Saccharomyces cerevisiae is a particularly attractive model because it displays pronounced cell polarity in response to intracellular and extracellular cues. Cells of S. cerevisiae undergo polarized growth during various phases of their life cycle, such as during vegetative growth, mating between haploid cells of opposite mating types, and filamentous growth upon deprivation of nutrition such as nitrogen. Substantial progress has been made in deciphering the molecular basis of cell polarity in budding yeast. In particular, it becomes increasingly clear how small GTPases regulate polarized cytoskeletal organization, cell wall assembly, and exocytosis at the molecular level and how these GTPases are regulated. In this review, we discuss the key signaling pathways that regulate cell polarization during the mitotic cell cycle and during mating.     

香菇135菌株特异SCAR标记在其原生质体单核中的分布

以香菇保藏菌种庆元135以及庆元出菇的135新鲜子实体组织分离得到的菌丝为材料制备原生质体,分别获得58和83个原生质体单核体;经交配型鉴定后,用已获得的135特异SCAR引物对单核进行PCR扩增,统计SCAR条带的分布。结果显示：原生质体单核体分为A1B1和A2B2两种交配型,而特异条带仅存在于后者中,证明135特异SCAR标记是稳定遗传的,也为鉴定以带标记核为亲本之一的杂交后代提供了依据,这就进一步拓宽了分子标记应用于菌株鉴定的适用范围。     

香菇135菌株特异SCAR标记在其原生质体单核中的分布

以香菇保藏菌种庆元135以及庆元出菇的135新鲜子实体组织分离得到的菌丝为材料制备原生质体,分别获得58和83个原生质体单核体;经交配型鉴定后,用已获得的135特异SCAR引物对单核进行PCR扩增,统计SCAR条带的分布。结果显示:原生质体单核体分为A1B1和A2B2两种交配型,而特异条带仅存在于后者中,证明135特异SCAR标记是稳定遗传的,也为鉴定以带标记核为亲本之一的杂交后代提供了依据,这就进一步拓宽了分子标记应用于菌株鉴定的适用范围。     

Preferential mating. Nonrandom mating of a continuous phenotype

A phenotypically determined nonrandom mating model is specified for a continuous trait controlled by a major gene. The distribution of phenotypic preferences of any particular individual choosing a mate may be specified in terms of the phenotype of the individual. The relative mating frequency is dependent on the frequency of the genotypes. Conditions for equilibria, polymorphic and degenerate, are given for a series of analytical and numerical cases. Certain classical nonrandom mating models for discrete phenotypes are special cases of the preferential mating model. However, other discrete phenotypic models do not have parallels because either the intergenotypic relationships are not geometrically consistent or the selective and assortative properties are not phenotypically consistent.     

Differential responses of yellow-breasted chats, Icteria virens, to male and female conspecific model presentations

A detailed understanding of the behavioural interactions between males and females is crucial for elucidating the selection pressures shaping mating system evolution; however, these interactions are often difficult to observe, particularly in free-living populations. We simulated extraterritorial intrusions by presenting conspecific models on to territories of a Neotropical migratory passerine, the yellow-breasted chat, Icteria virens. Simulated intrusions elicited different responses based on the sex of the focal individual and the sex and species of the model. Behavioural responses of the focal individuals were largely directed towards conspecific models, whereas simulated heterospecific intrusions, using a Carolina wren, Thryothorus ludovicianus, model, elicited minimal response. Male song was not related to any particular model presentation. Males directed mating behaviours, including courtship displays and copulations, exclusively towards the female conspecific model. Males behaved aggressively towards the conspecific male model. In contrast, females never showed any courtship behaviour towards the male model and showed significant aggression towards the conspecific female model. The results of this study reveal that territorial aggression in yellow-breasted chats is strongly intrasexual. Additionally, simulated female extraterritorial intrusions, but not male intrusions, resulted in extrapair courtship and copulation. This pattern is likely to be typical of many socially monogamous species and points to the behavioural mechanisms underlying both monogamy and extrapair mating systems.     

Mapping the mating type locus of Tetrahymena thermophila: meiotic linkage of mat to the ribosomal RNA gene.

Tetrahymena thermophila has a multiple mating type system. While a sexually mature cell usually expresses only one mating type, its germline (micronucleus) carries the genetic potential for 5 to 7 mating types. The set of allowed mating types is specified by the mat locus. The choice of which particular mating type is expressed by a cell reflects a somatically inherited, developmentally programmed differentiation of the somatic nucleus (macronucleus). In this work we report that the mat locus maps to the left arm of chromosome 2, as determined by nullisomic deletion mapping. We also report a distance of 29 cM between the mat locus and the ribosomal RNA gene, previously mapped to chromosome 2L. This represents another (rare) case of meiotic linkage in Tetrahymena.     

Cockroach mating behaviors,sex pheromones,and abdominal glands (Dictyoptera: Blaberidae)

Two chemical signals are essential in all cockroach sexual behavioral sequences: the sex pheromone released by one partner, generally the female (for long distance attraction), and an aphrodisiac sex pheromone produced exclusively by male tergal glands (for female mounting and tergal contact or feeding behavior). Unlike the other cockroach groups, the males of the Oxyhaloinae species produce both chemical signals: the pheromone and the aphrodisiac. The occurrence of three patterns of mating behavior (A, B, and C), the production of male sex pheromones, and the existence in the male of developed sternal and tergal glands in seven related Oxyhaloinae species, make these cockroaches a useful model for studying the evolution of mating behavior patterns. The various types of mating behavior were not classified in the previous studies by Roth and Barth. In this report, they have been named type A (female in upper position), B (male in upper position), and C (male and female end to end). In type A mating, the male tergal glands, which are licked by the females, are well developed, whereas in types B and C, there is no licking of the male's tergal secretion by the females and the tergal glands are much less developed; the aphrodisiacs secreted by the tergal glands may no longer act in this case through contact chemoreception, but through an olfactory process involving volatile components. One common sex pheromone component seems to be acetoin. I suggest that the mating behavior tends from A toward B and C during the evolutionary process with a concomitant regression of the tergal glands and changes in the aphrodisiac emission levels. The mating behavioral sequences of cockroaches (Dictyoptera) and crickets (Orthoptera) show a striking degree of similarity and are probably examples of convergent evolution.     

Reinforcement and the genetics of nonrandom mating

Abstract.— The occurrence of reinforcement is compared when premating isolation is caused by the spread of a gene causing females to prefer to mate with males carrying a population-specific trait (a "preference" model) and by a gene that causes females to prefer to mate with males that share their own trait phenotype (an "assortative mating" model). Both two-island models, which have symmetric gene flow, and continent-island models, which have one-way gene flow, are explored. Reinforcement is found to occur much more easily in a two-island assortative mating model than in any of the other three models. This is due primarily to the fact that in this model the assortative mating allele will automatically become genetically associated in each population with the trait allele that is favored by natural selection on that island. In contrast, natural selection on the trait both favors and opposes the evolution of premating isolation in the two-island preference model, depending on the particular population. These results imply that species recognition in the context of mating may evolve particularly easily when it targets cues that are favored by natural selection in each population. In the continent-island models, reinforcement is found to occur more often under the preference model than the assortative mating model, thus reversing the trend from the two-island models. Patterns of population subdivision may therefore play a role in determining what types of premating isolation may evolve.     

Ecological Genetics of Tetrahymena thermophila: Mating Types, i-Antigens, Multiple Alleles and Epistasis

Until recently, Tetrahymena thermophila has rarely been isolated from nature. With improved sampling procedures, T. thermophila has been found in ponds in many northeastern states. The availability of resident populations makes possible both population and ecological genetic studies. All seven known mating types have been recovered; no eighth mating type has been found. Crosses among whole-genome homozygotes derived from Pennsylvania isolates reveal a spectrum genotypes with mating type alleles resembling traditional A (IV- and VII-) and B(I-) categories. The genotypes differ significantly with respect to mating type frequency, both among themselves and from previously described genotypes. One A-category genotype appears to lack mating type II, while one A-category and all B-category genotypes have low frequencies of mating type III, thus accounting for the low frequency of III in the pond. The low frequency of III in all five B-category genotypes examined suggests that the founding allele in this region was low for III. These and other differences are discussed both in terms of mating type frequencies in the pond and in terms of the possible molecular structure of mat alleles. By contrast, numerous variants of the cell surface immobilization antigen are found in addition to the previously described i-antigens. Variants of the known SerH alleles include those with restriction fragment length polymorphisms and temperature sensitivity as well as alleles with new antigenic specificity. Multiple alleles are present in single ponds. Genes exhibiting serially dominant epistasis over SerH genes also are found. In two instances (K and C), families of antigenically similar polypeptides are expressed in place of H i-antigen. Molecular weight differences suggest that these paralogous i-antigen genes evolve by gene duplication and unequal crossing over within central repeats. The existence of complex patterns of epistasis together with seasonal changes in i-ag frequencies suggest that i-ag play an important, but as yet unknown, ecological role related to the occurrence of frequent conjugation.     

Evolution of hierarchical cytoplasmic inheritance in the plasmodial slime mold Physarum polycephalum

A striking linear dominance relationship for uniparental mitochondrial transmission is known between many mating types of plasmodial slime mold Physarum polycephalum. We herein examine how such hierarchical cytoplasmic inheritance evolves in isogamous organisms with many self-incompatible mating types. We assume that a nuclear locus determines the mating type of gametes and that another nuclear locus controls the digestion of mitochondria DNAs (mtDNAs) of the recipient gamete after fusion. We then examine the coupled genetic dynamics for the evolution of self-incompatible mating types and biased mitochondrial transmission between them. In Physarum, a multiallelic nuclear locus matA controls both the mating type of the gametes and the selective elimination of the mtDNA in the zygotes. We theoretically examine two potential mechanisms that might be responsible for the preferential digestion of mitochondria in the zygote. In the first model, the preferential digestion of mitochondria is assumed to be the outcome of differential expression levels of a suppressor gene carried by each gamete (suppression-power model). In the second model (site-specific nuclease model), the digestion of mtDNAs is assumed to be due to their cleavage by a site-specific nuclease that cuts the mtDNA at unmethylated recognition sites. Also assumed is that the mtDNAs are methylated at the same recognition site prior to the fusion, thereby being protected against the nuclease of the same gamete, and that the suppressor alleles convey information for the recognition sequences of nuclease and methylase. In both models, we found that a linear dominance hierarchy evolves as a consequence of the buildup of a strong linkage disequilibrium between the mating-type locus and the suppressor locus, though it fails to evolve if the recombination rate between the two loci is larger than a threshold. This threshold recombination rate depends on the number of mating types and the degree of fitness reduction in the heteroplasmic zygotes. If the recombination rate is above the threshold, suppressor alleles are equally distributed in each mating type at evolutionary equilibrium. Based on the theoretical results of the site-specific nuclease model, we propose that a nested subsequence structure in the recognition sequence should underlie the linear dominance hierarchy of mitochondrial transmission.     

More than just noise: Chance,mating success,and sexual selection

Chance plays a critical but underappreciated role in determining mating success. In many cases, we tend to think of chance as background noise that can be ignored in studies of mating dynamics. When the influence of chance is consistent across contexts, chance can be thought of as background noise; in other cases, however, the impact of chance on mating success can influence our understanding of how mates are acquired and how sexual selection operates. In particular, when the importance of chance covaries with biological or ecological factors in a systematic manner—that is, when chance becomes consistently more or less important under certain conditions—then chance is important to consider if we want to fully understand the operation of mate acquisition and sexual selection. Here, we present a model that explores how chance covaries with factors such as sex ratio, adult population size, and mating regime in determining variation in mating success. We find that in some cases, chance covaries with adult population size and the operational sex ratio to create variation in mating success. We discuss how chance can influence our more general understanding of the operation of mating dynamics and sexual selection.