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PubMed1986

The significance of over- and underdominance for the maintenance of genetic polymorphisms. II. Overdominance and instability with random mating.

Part I of the present series demonstrates that globally stable polymorphic equilibria may show underdominance in Darwinian fitness. Hence, overdominance in fitness can no longer be conceived of as a necessary condition for the stability of a polymorphism. In the present paper, the question is posed as to whether overdominance is at least sufficient for this stability. A population of randomly mating individuals is considered, where selection operates uniquely through differential fecundities of particular mating types and may generate either a heterozygote excess or deficit relative to Hardy-Weinberg proportions. It turns out that both unstable central overdominance and stable central underdominance are possible and that their occurrence is strongly related to an excess or a deficiency of heterozygotes in the vicinity of the regions of instability or stability. As one consequence, the above suggested sufficiency of heterozygote superiority is not valid, even in random mating populations. Based on the results of both papers of this series, which demonstrate the inadequacy of over- and underdominance as indicators of stability or instability, a modified overdominance principle is discussed. This principle states that a biallelic polymorphism is maintained if the heterozygote is superior in its degree of "heterogamous self-replication" to the degrees of "autogamous self-replication" of the corresponding homozygotes. It is derived with the help of fractional fitnesses, and it is pointed out that certain ratios of these may be more useful for finding evolutionary constants which govern the maintenance of genetic polymorphisms than are ratios of total fitnesses.

Animals↗
PubMed2004

Sexual selection can constrain sympatric speciation.

Recent theory has suggested that sympatric speciation can occur quite easily when individuals that are ecologically similar mate assortatively. Although many of these models have assumed that individuals have equal mating success, in nature rare phenotypes may often suffer decreased mating success. Consequently, assortative mating may often generate stabilizing sexual selection. We show that this effect can substantially impede sympatric speciation. Our results emphasize the need for data on the strength of the stabilizing component of selection generated by mating in natural populations.

Animals↗
PubMed1996

Yeast alpha mating factor structure-activity relationship derived from genetically selected peptide agonists and antagonists of Ste2p.

alpha-Factor, a 13-amino-acid pheromone secreted by haploid alpha cells of Saccharomyces cerevisiae, binds to Ste2p, a seven-transmembrane, G-protein-coupled receptor present on haploid alpha cells, to activate a signal transduction pathway required for conjugation and mating. To determine the structural requirements for alpha-factor activity, we developed a genetic screen to identify from random and semirandom libraries novel peptides that function as agonists or antagonists of Ste2p. The selection scheme was based on autocrine strains constructed to secrete random peptides and respond by growth to those that were either agonists or antagonists of Ste2p. Analysis of a number of peptides obtained by this selection procedure indicates that Trp1, Trp3, Pro8, and Gly9 are important for agonist activity specifically. His2, Leu4, Leu6, Pro10, a hydrophobic residue 12, and an aromatic residue 13 are important for both agonist and antagonist activity. Our results also show that activation of Ste2p can be achieved with novel, unanticipated combinations of amino acids. Finally, the results suggest the utility of this selection scheme for identifying novel ligands for mammalian G-protein-coupled receptors heterologously expressed in S. cerevisiae.

Amino Acid Sequence↗
PubMed2006

Good genes sexual selection in nature.

Whether the mate sampling and choice performed by females in nature influences offspring performance is a controversial issue in theory and an open empirical question. Pronghorn (Antilocapra americana) females engage in an obvious and energetically expensive mate sampling process to identify vigorous males. Although individual females sample independently, their choices converge on a small proportion of males that sire most young. Offspring of attractive males were more likely to survive to weaning and to age classes as late as 5 years, resulting in a selection differential, calculated by expected differences in lifetime number of offspring weaned, of 0.32 against random mating. Enhanced survival to weaning appeared to be accomplished by faster growth rates. Females compensated for matings with a less attractive mate by elevating rates of milk delivery to their young. Because pronghorn males do not have costly ornaments, we conclude that female choice for good genes can exist in the absence of ornaments. Furthermore, female choice may be important and unrecognized as a force that can lower population genetic load.

Animals↗
PubMed2005

Mate choice decisions of stickleback females predictably modified by MHC peptide ligands.

Sexual selection has been proposed as one mechanism to explain the maintenance of high allelic diversity in MHC genes that control the extent of resistance against pathogens and parasites in natural populations. MHC-based sexual selection is known to involve olfactory mechanisms in fish, mice, and humans. During mate choice, females of the three-spined stickleback (Gasterosteus aculeatus) use an odor-based selection strategy to achieve an optimal level of MHC diversity in their offspring, equipping them with optimal resistance toward pathogens and parasites. The molecular mechanism of odor-based mate-selection strategies is unknown. Because peptide ligands for MHC class I molecules function as individuality signals in mice, we hypothesized that female sticklebacks might assess the degree of MHC diversity of potential partners by means of the structural diversity of the corresponding peptide ligands in perceived odor signals. We show that structurally diverse MHC ligands interact with natural odors of male sticklebacks to predictably modify MHC-related mate choice. For a mating pair with suboptimal numbers of MHC alleles, peptides increase the attractiveness of male water, whereas for a mating pair with superoptimal numbers, attractiveness is decreased. Our results suggest that female sticklebacks use evolutionarily conserved structural features of MHC peptide ligands to evaluate MHC diversity of their prospective mating partners.

Amino Acid Sequence↗
PubMed2005

Speciation through the learning of habitat features.

Learning of environmental features can influence both mating behaviour and the location where young are produced. This may lead to speciation in three steps: (i) colonization of a new habitat, (ii) genetic divergence of the two groups by adaptation to the habitats, and (iii) a decrease of genetic mixing between the lineages (similar to reinforcement). In a previous paper we showed that steps (i) and (ii) occur readily for a wide range of fixed mating and habitat preferences. Here, we study whether this can ultimately lead to speciation through selective changes in these preferences. We show that this indeed occurs, and, furthermore, it is very general: for a large class of models there is selection toward producing young more frequently in the natal habitat. Once habitat preference is strong, there is selection toward stronger assortative mating. Even when steps (i) and (ii) initially fail, genetic divergence may succeed at a later evolutionary stage, after which a decrease of genetic mixing completes speciation. Our results show that speciation by the learning of habitat features is an extremely effective mechanism.

Animals↗
PubMed1990

Calving disorders of primiparous Holsteins from designed selection studies.

Primiparous Holsteins from designed studies were evaluated for edema, dystocia, and retained placenta. Genetic studies were 1) selection for large body size versus small body size, and 2) milk yield selection versus control (no selection). The large body size group was mated to sires with extreme estimates of transmitting ability for tall height and deep and wide bodies. Small body size group was mated to extreme sires for transmitting short height and shallow, narrow bodies. Large and small cows significantly differed only for edema, and small cows had scores suggesting more severe udder edema than large cows. Milk selection group was mated to the four highest sires for PTA milk each year, whereas a control group was mated to 20 randomly selected AI sires that were breed average for PTA milk in 1964. Selection groups for milk yield did not differ significantly for any of the calving disorders.

Age Factors↗
PubMed2006

Complex multivariate sexual selection on male acoustic signaling in a wild population of Teleogryllus commodus.

Mate choice may impose both linear (i.e., directional) and nonlinear (i.e., quadratic and correlational) sexual selection on advertisement traits. Traditionally, mate recognition and sensory tuning have been thought to impose stabilizing (i.e., negative quadratic) sexual selection, whereas adaptive mate choice effects directional selection. It has been suggested that adaptive choice may exert positive quadratic and/or correlational sexual selection. Earlier, we showed that five structural components of the advertisement call of male field crickets (Teleogryllus commodus) were under multivariate stabilizing selection under laboratory conditions. Here we experimentally estimate selection on these five traits plus a measure of calling activity (the number of repeats in a looped bout of calling) in the field. There was general support for multivariate stabilizing selection on call structure, and calling activity was under strong positive directional selection, as predicted for a signal of genetic quality. There was, however, also appreciable correlational selection, suggesting an interaction between male call structure and calling effort. Interestingly, selection for short interbout durations of silence favored longer intercall durations in the field, in contrast to results from continuous looped call playback in the laboratory. We discuss the general importance of nonlinear selection in the honest signaling of genetic quality.

Acoustic Stimulation↗
PubMedDate not supplied

Testing alternative methods for purging genetic load using the housefly (Musca domestica L.).

When a population faces long-term inbreeding, artificial selection, in principle, can enhance natural selection processes for purging the exposed genetic load. However, strong purge pressures might actually decrease fitness through the inadvertent fixation of deleterious alleles and allelic combinations. We tested lines of the housefly (Musca domestica L.) for the effectiveness of artificial selection to promote the adaptation to small population size. Specifically, replicate populations were held at average census sizes of 54 for nine generations or 30 for 14 generations while being subjected to artificial selection pressure for increased fitness in overall mating propensity (i.e., the proportion of virgin male-female pairs initiating copulation within 30 min), while also undergoing selection to create differences among lines in multivariate components of courtship performance. In the 14-generation experiment, a subset of the lines were derived from a founder-flush population (i.e., derived from three male-female pairs). In both experiments, we also maintained parallel non-selection lines to assess the potential for natural purging through serial inbreeding alone. Sub-populations derived from a stock newly derived from the wild responded to artificial selection for increased mating propensity, but only in the short-term, with eventual rebounds back to the original levels. Serial inbreeding in these lines simply reduced mating propensity. In sub-populations derived from the same base population, but 36 generations later, both artificial selection and serial inbreeding increased mating propensity, but mainly to restore the level found upon establishment in the laboratory. Founder-flush lines responded as well as the non-bottlenecked controls, so we base our major conclusions on the comparisons between fresh-caught and long-term laboratory stocks. We suggest that the effectiveness of the alternative purge protocols depended upon the amount of genetic load already exposed, such that prolonged periods of relaxed or altered selection pressures of the laboratory rendered a population more responsive to purging protocols.

Animals↗
PubMed1994

Sex differences, sex ratios and sex roles.

Sexual selection theory predicts that sex roles will be determined by the operational sex ratio (OSR), the sex ratio among individuals searching for mates at any given time. There are two predictions: (i) the sex which is in 'excess' will be the more competitive sex with respect to access to mates; and (ii) the sex of which there is a 'shortage' will be the more choosy with respect to potential partners. We examine the second prediction and find that current OSR theory does not consider an important factor which affects mate choice. This factor is sex differences in variation in mate quality. Hence, we develop a new model of mate choice which shows that the parameter which should be optimized during mate choice is the trade-off between reproductive rate and mate quality. If mate choice is too lax, reproductive rate may be high but partners will be of low quality. If mate choice is too stringent, partners will be of high quality but reproductive rate will be low because such partners will be rare. Stringency of mate choice is, therefore, a facet of OSR theory. Indeed, our model shows that OSR theory can be used to integrate the effect of sex differences in both mating rate and variation in mate quality to predict the direction of mate choice. Our model suggests that: (i) mate choice is only selected when individuals of the opposite sex vary in their quality as mates; (ii) if the extent of variation in mate quality is equal within each sex, the sex with the lower potential mating rate will be the more choosy sex; but (iii) if there is sufficiently greater variation in mate quality among the sex with the lower potential reproductive rate, the sex with the higher potential mating rate will be the more choosy sex.(ABSTRACT TRUNCATED AT 250 WORDS)

Game Theory↗
PubMed1975

Sex differences in gene expression for pupa weight in long term selected lines of Tribolium.

Full-sib matings in two populations of Tribolium which had been selected for increased pupa weight for more than 85 generations resulted in a significant inbreeding depression in male progeny but showed no effect in the female progeny. An analysis of variance of a population produced by backcrossing the selected populations to the inbred lines originally used to establish the select populations (Design-III) indicated some genes were still segregating which produce dominance effects in males but not in females. The data support the hypothesis that a class of genes exists, associated with the autosomes, that differ in their dominance effects of pupa weight in the two sexes.

Animals↗
PubMed1999

Effect of selection against deleterious mutations on the decline in heterozygosity at neutral loci in closely inbreeding populations.

Transition matrices for selfing and full-sib mating were derived to investigate the effect of selection against deleterious mutations on the process of inbreeding at a linked neutral locus. Selection was allowed to act within lines only (selection type I) or equally within and between lines (type II). For selfing lines under selection type I, inbreeding is always retarded, the retardation being determined by the recombination fraction between the neutral and selected loci and the inbreeding depression from the selected locus, irrespective of the selection coefficient (s) and dominance coefficient (h) of the mutant allele. For selfing under selection type II or full-sib mating under both selection types, inbreeding is delayed by weak selection (small s and sh), due to the associative overdominance created at the neutral locus, and accelerated by strong selection, due to the elevated differential contributions between alternative alleles at the neutral locus within individuals and between lines (for selection type II). For multiple fitness loci under selection, stochastic simulations were run for populations with selfing, full-sib mating, and random mating, using empirical estimates of mutation parameters and inbreeding load in Drosophila. The simulations results are in general compatible with empirical observations.

Animals↗
PubMed1993

Honest signalling among gametes.

The gametes of many lower eukaryotic organisms emit pheromones that attract gametes of the opposite mating type or sex. Gametes move or grow in the direction of the highest pheromone concentration, suggesting that the strength of the pheromonal signal is used to infer proximity, or that the strongest signal is most likely to be notice. Here I offer a new explanation of pheromonal signalling and chemotaxis in gametes. I show that pheromonal signals can be interpreted as sexually selected traits that honestly advertise variation in quality among gametes, given that signals are costly to produce and that gametes compete; by 'quality' I refer to some aspect of a gamete's fitness. A gamete's preference for a mating partner, then, is predicted to vary with the quality of a prospective partner as inferred from the strength of its signal. This view can explain characteristics of the signalling and mate selection behaviours of gametes that are not predicted by models of mate choice based on proximity or 'passive attraction' to the strongest signal. These include repeated partner exchanges, escalated exchanges of mating pheromones, and rejection of gametes that signal at low levels.

Animals↗
PubMed2003

Sexual dimorphism and adaptive speciation: two sides of the same ecological coin.

Models of adaptive speciation are typically concerned with demonstrating that it is possible for ecologically driven disruptive selection to lead to the evolution of assortative mating and hence speciation. However, disruptive selection could also lead to other forms of evolutionary diversification, including ecological sexual dimorphisms. Using a model of frequency-dependent intraspecific competition, we show analytically that adaptive speciation and dimorphism require identical ecological conditions. Numerical simulations of individual-based models show that a single ecological model can produce either evolutionary outcome, depending on the genetic independence of male and female traits and the potential strength of assortative mating. Speciation is inhibited when the genetic basis of male and female ecological traits allows the sexes to diverge substantially. This is because sexual dimorphism, which can evolve quickly, can eliminate the frequency-dependent disruptive selection that would have provided the impetus for speciation. Conversely, populations with strong assortative mating based on ecological traits are less likely to evolve a sexual dimorphism because females cannot simultaneously prefer males more similar to themselves while still allowing the males to diverge. This conflict between speciation and dimorphism can be circumvented in two ways. First, we find a novel form of speciation via negative assortative mating, leading to two dimorphic daughter species. Second, if assortative mating is based on a neutral marker trait, trophic dimorphism and speciation by positive assortative mating can occur simultaneously. We conclude that while adaptive speciation and ecological sexual dimorphism may occur simultaneously, allowing for sexual dimorphism restricts the likelihood of adaptive speciation. Thus, it is important to recognize that disruptive selection due to frequency-dependent interactions can lead to more than one form of adaptive splitting.

Adaptation, Biological↗
PubMed2003

Cuticular pheromones and water balance in the house fly, Musca domestica.

Epicuticular lipids serve two major roles in insects. Their waterproofing properties are crucial to survival in terrestrial environments, and they serve as contact pheromones in a wide array of taxa. Both functions may be affected by the physical properties of the surface lipids. This provides the opportunity for natural selection on water conservation, mediated by lipid phase behavior, to interact with and perhaps conflict with sexual selection on communication and mate recognition. We used the common house fly, Musca domestica, as a model for these interacting selective forces. Male house flies preferred female models treated with a high melting-point lipid mixture, suggesting that sexual and natural selection may both act to favor longer-chain, more saturated hydrocarbons. However, higher melting points did not result in lower rates of water loss. We propose a working model in which phase separation between the unsaturated female pheromone and saturated hydrocarbons results in areas of melted, pheromone-rich lipids and regional variation in cuticular permeability.

Age Factors↗