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Exploring the consequences of postmating-prezygotic interactions between the sexes.

The role of male-female interactions in the divergence of postmating-prezygotic reproductive traits has recently focused on sexual conflict as the selective force. While an association between mating costs and benefits suggests that antagonistic interactions may be important, a mosaic of processes may actually mediate the evolutionary dynamics of postmating-prezygotic interactions. Our study examines the ecological relevance of mating effects on females. We test this critical but often overlooked assumption in desert Drosophila, where the species' ecology provides a framework for evaluating the evolutionary implications of such mating consequences. We show that mating has a profound impact on an ecologically critical character--desiccation resistance. To explore what selective factors might underlie the observed population differences in mating effects, we test whether trends in the data match predictions based on the species' ecology. While these preliminary data are consistent with the expectation that the mating benefits are positively correlated with environmental conditions, further examination reveals an additional unanticipated association with the reproductive environment. This unexpected association challenges existing assumptions regarding the forces driving divergence of traits involved in mating.

Animals↗

Post-mating clutch piracy in an amphibian.

Female multiple mating and alternative mating systems can decrease the opportunity for sexual selection. Sperm competition is often the outcome of females mating with multiple males and has been observed in many animals, and alternative reproductive systems are widespread among species with external fertilization and parental care. Multiple paternity without associated complex behaviour related to mating or parental care is also seen in simultaneously spawning amphibians and fishes that release gametes into water. Here we report 'clutch piracy' in a montane population of the common frog Rana temporaria, a reproductive behaviour previously unknown in vertebrates with external fertilization. Males of this species clasp the females and the pair deposits one spherical clutch of eggs. No parental care is provided. 'Pirate' males search for freshly laid clutches, clasp them as they would do a female and fertilize the eggs that were left unfertilized by the 'parental' male. This behaviour does not seem to be size-dependent, and some males mate with a female and perform clutch piracy in the same season. Piracy affected 84% of the clutches and in some cases increased the proportion of eggs fertilized, providing direct fitness benefits both for the pirate males and the females. Sexual selection--probably caused by a strong male-biased sex ratio--occurs in this population, as indicated by size-assortative mating; however, clutch piracy may reduce its impact. This provides a good model to explore how alternative mating strategies can affect the intensity of sexual selection.

Animals↗

Direct and correlated responses to selection for milk yield: results and conclusions of regional project NC-2, "improvement of dairy cattle through breeding, with emphasis on selection". NC-2 Technical Committee.

Measurement of direct and correlated responses to single-trait selection for milk yield was the major objective of regional project NC-2. The NC-2 Technical Committee included representatives from Alaska, Illinois, Indiana, Iowa, Kansas, Michigan, Minnesota, Nebraska, South Dakota, Wisconsin, and the USDA. All representatives, except Illinois, Kansas and Nebraska, maintained a selection line formed by using AI sires selected for high estimated transmitting abilities for milk and a second line that served as some type of a control. Stations varied in criteria for selection of bulls for control lines. Farms were managed similarly, including feeding and management of selection and control lines as one herd, random mating within line, and restricted culling policies. Selection for milk yield effectively increased milk production. All selection lines increased milk and net income per lactation more than control lines. Realized gains matched or exceeded gains expected from estimates of breeding values. Yields of milk components increased, but component percentages decreased appreciably for selection lines. Reproduction of nulliparous animals was not affected, but days open for lactating selection cows increased in some of the individual projects. Selected cows tended to have larger health costs, specifically for mammary treatment. Udder and conformation traits did not deteriorate for selection lines, although control lines with selection of sires on genetic evaluations for type received higher type scores. There should be few reservations about undesirable responses correlated with selection for milk yield.

Animal Feed↗

Adaptive speciation when assortative mating is based on female preference for male marker traits.

Adaptive speciation occurs when frequency-dependent ecological interactions generate conditions of disruptive selection to which lineage splitting is an adaptive response. Under such selective conditions, evolution of assortative mating mechanisms enables the break-up of the ancestral lineage into diverging and reproductively isolated descendent species. Extending previous studies, I investigate models of adaptive speciation due to the evolution of indirect assortative mating that is based on three different mating traits: the degree of assortativity, a female preference trait and a male marker trait. For speciation to occur, linkage disequilibria between different mating traits, e.g. between female preference and male marker traits, as well as between mating traits and the ecological trait, must evolve. This can lead to novel speciation scenarios, e.g. when reproductive isolation is generated by a splitting in the degree of assortativeness, with one of the emerging lineages mating assortatively, and the other one disassortatively. I investigate the effects of variation in various model parameters on the likelihood of speciation, as well as robustness of speciation to introducing costs of assortative mating. Even though in the models presented speciation requires the genetic potential for strong assortment as well as rather restrictive ecological conditions, the results show that adaptive speciation due to the evolution of assortative mating when mate choice is based on separate female preference and male marker traits is a theoretically plausible evolutionary scenario.

Adaptation, Biological↗

Experimental evolution of mating discrimination in budding yeast.

Assortative mating, when individuals of similar phenotypes mate, likely plays a key role in preventing gene flow during speciation. Reinforcement occurs when two previously geographically separated (allopatric) groups meet after having evolved partial postzygotic isolation; they are selected to evolve or enhance assortative mating to prevent costly intergroup matings that produce only maladaptive or sterile hybrids. Studies in Drosophila have shown that the genetic architectures of mating discrimination could differ significantly with or without reinforcement, suggesting that the evolution of assortative mating may be more complicated than expected. To study the evolution of assortative mating, we evolved mating discrimination in populations of the budding yeast, Saccharomyces cerevisiae. After 36 cycles of selection, these cells are five times more likely to mate with each other than to their ancestors, despite detectable one-way gene flow between the selected and reference populations. Several individual cultures evolved mating discrimination by changing their mating kinetics, with some mating more rapidly and others more slowly than the ancestral population. Genetic analysis indicates that multiple mutations have accumulated to produce the altered mating preference. Our results show that subtle details of mating behavior can play an important role in the evolution of reproductive isolation.

Biological Evolution↗

Lifetime mating opportunities and male mating behaviour in sexually cannibalistic praying mantids.

I examined the number of lifetime mating opportunities and mating behaviour of males in two sexually cannibalistic species, the Mediterranean, Iris oratoria, and bordered, Stagmomantis limbata, praying mantids (Mantodea: Mantidae). Two approaches estimated the number of lifetime mating opportunities: direct observations of intersexual encounters in the field, and an encounter model. I collected behavioural observations, together with ecological data for use in the model, over three field seasons. The ecological data included an assessment of the feeding condition of S. limbata females in nature; the females fed at a level comparable to females maintained on an abundant diet in the laboratory. As for the number of mating opportunities, individual males of both species encountered two or more females, as predicted by the model. I observed no male, however, in more than one copulation. This result could reflect individual variation in the times and places of sexual activity or an actual low number of mating opportunities in the field. Furthermore, a higher percentage of I. oratoria males encountered two or more females than S. limbata males, as the model indicates. Fewer mating opportunities could lead to greater selection upon S. limbata males to ensure paternity at each mating, which can explain the longer copulation times observed for S. limbata males. I considered two hypotheses about male behaviour in light of the number of lifetime encounters with females: male suicide and male reduction of the risk of cannibalism. Behavioural observations do not strongly support male suicide in either species. Certain male behaviours, such as the nature of copulatory position and, in captivity, mounting females from the rear, are consistent with the idea that males behave so as to reduce the probability that they are cannibalized during intersexual encounters. Moreover, male I. oratoria preferentially mount well-fed, fecund females in captivity. Taken together, these results suggest that the prospect of multiple mating opportunities has influenced male mating behaviour over evolutionary time. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Mitochondria are inherited from the MATa parent in crosses of the basidiomycete fungus Cryptococcus neoformans.

Previous studies demonstrated that mitochondrial DNA (mtDNA) was uniparentally transmitted in laboratory crosses of the pathogenic yeast Cryptococcus neoformans. To begin understanding the mechanisms, this study examined the potential role of the mating-type locus on mtDNA inheritance in C. neoformans. Using existing isogenic strains (JEC20 and JEC21) that differed only at the mating-type locus and a clinical strain (CDC46) that possessed a mitochondrial genotype different from JEC20 and JEC21, we constructed strains that differed only in mating type and mitochondrial genotype. These strains were then crossed to produce hyphae and sexual spores. Among the 206 single spores analyzed from six crosses, all but one inherited mtDNA from the MATa parents. Analyses of mating-type alleles and mtDNA genotypes of natural hybrids from clinical and natural samples were consistent with the hypothesis that mtDNA is inherited from the MATa parent in C. neoformans. To distinguish two potential mechanisms, we obtained a pair of isogenic strains with different mating-type alleles, mtDNA types, and auxotrophic markers. Diploid cells from mating between these two strains were selected and 29 independent colonies were genotyped. These cells did not go through the hyphal stage or the meiotic process. All 29 colonies contained mtDNA from the MATa parent. Because no filamentation, meiosis, or spore formation was involved in generating these diploid cells, our results suggest a selective elimination of mtDNA from the MATalpha parent soon after mating. To our knowledge, this is the first demonstration that mating type controls mtDNA inheritance in fungi.

Crosses, Genetic↗

Transferable drug resistance in Pseudomonas aeruginosa.

Three strains of Pseudomonas aeruginosa were demonstrated to transfer double-drug resistance by conjugation to a P. aeruginosa recipient at frequencies of 10(-4) to 10(-2) per recipient cell. Two of the three strains also transferred to Escherichia coli at frequencies which were 10(3)- to 10(5)-fold lower, but the third strain could not be demonstrated to do so. The latter strain, however, conferred maleness on the Pseudomonas recipient. The transfer of streptomycin resistance was associated with the acquisition of streptomycin phosphorylase by both P. aeruginosa and E. coli recipients. Maximal broth mating frequencies were obtained with nonagitated cultures less than 1 mm in depth. A pyocine selection system based on donor sensitivity and recipient resistance is described and appears to have future value as a generalized selective device for use after matings.

Conjugation, Genetic↗

Genetic exchange of transposon and integrative plasmid markers in Mycoplasma pulmonis.

Matings of genetically marked derivatives of Mycoplasma pulmonis resulted in the exchange of chromosomal DNA and the appearance of doubly marked transconjugants. Transposons Tn916 and Tn4001, and a series of integrative plasmids derived from their cloned antibiotic resistance genes, were used to construct antibiotic-resistant mycoplasmal derivatives to examine this phenomenon at the molecular level. Genetic exchange occurred on agar surfaces at frequencies ranging from 3.3 X 10(-4) to 6.4 X 10(-8) transconjugants per CFU. Examination of chromosomal DNA from transconjugants by hybridization revealed that the transposons or integrated plasmids were in the same chromosomal locations as in the parental strains, indicating that exchange involved the transfer of chromosomal DNA and homologous recombination. Transfer was not affected by DNase, polyethylene glycol, EDTA, or calcium chloride but was affected by treatment of either parent with trypsin. Mixing of mating strains before plating had no effect on mating frequencies, but mating did occur in liquid media. The ability to exchange chromosomal markers was limited to selected strains of M. pulmonis; mating did not occur with Acholeplasma laidlawii or M. gallisepticum. Heat and UV inactivation studies revealed that nonviable cells could act as donors in matings. The evidence presented supports a conjugationlike mechanism involving specific trypsin-sensitive membrane components.

Acholeplasma laidlawii↗

Genetic variation for West Nile virus susceptibility in Culex tritaeniorhynchus.

Mutant and geographical strains of Culex tritaeniorhynchus were compared for West Nile (WN) virus susceptibility by feeding on a high-titered blood-virus suspension. Eleven strains also were selected from 2-21 generations for an increase and/or a reduction of oral susceptibility using 90% and 10% infective virus doses, respectively. Only one of the 20 strains tested, e ma, was significantly less susceptible than the control strain. In the selection experiments, none of the strains showed a consistent decrease in susceptibility, but the Changa Manga II (CM) strain showed a sustained increase in susceptibility from generations F11-F21 when selection was discontinued. Reciprocal cross-matings and back-crosses were set up between the selected CM strain and two of the morphological mutant strains, e ma and re e ae, carrying homozygous recessive markers. The resulting progeny were tested for susceptibility to WN virus infection and the ability to replicate virus to high-titers after infection. These results suggest that the trait of increased susceptibility is dominant over resistance. The enhanced ability of infected mosquitoes to replicate WN virus showed partial dominance. Both of these traits appear to be polyfactorial, and are apparently associated with more than one chromosome in Cx. tritaeniorhynchus.

Animals↗

The evolution of male traits in social insects.

Pair formation in social insects mostly happens early in adult life and away from the social colony context, which precludes promiscuity in the usual sense. Termite males have continuous sperm production, but males of social Hymenoptera have fixed complements of sperm, except for a few species that mate before female dispersal and show male-fighting and lifelong sperm production. We develop an evolutionary framework for testing sexual selection and sperm competition theory across the advanced eusocial insects (ants, wasps, bees, termites) and highlight two areas related to premating sexual selection (sexual dimorphism and male mate number) that have remained understudied and in which considerable progress can be achieved with relatively simple approaches. We also infer that mating plugs may be relatively common, and we review further possibilities for postmating sexual selection, which gradually become less likely in termite evolution, but for which eusocial Hymenoptera provide unusual opportunities because they have clonal ejaculates and store viable sperm for up to several decades.

Animals↗

Seasonal cycles in the blood plasma concentration of FSH, inhibin and testosterone, and testicular size in rams of wild, feral and domesticated breeds of sheep.

Seasonal cycles in testicular activity in rams were monitored in groups of wild (mouflon), feral (Soay) and domesticated breeds of sheep (Shetland, Blackface, Herdwick, Norfolk, Wiltshire, Portland and Merino) living outdoors near Edinburgh (56 degrees N). The changes in the blood plasma concentrations of FSH, inhibin and testosterone, and the diameter of the testis were measured every half calendar month from 1 to 3 years of age. There were significant differences between breeds in the magnitude and timing of the seasonal reproductive cycle. In the mouflon rams, the seasonal changes were very pronounced with a 6-15-fold increase in the plasma concentrations of FSH, inhibin and testosterone from summer to autumn, and a late peak in testicular diameter in October. In the Soay rams and most of the domesticated breeds, the seasonal increase in the reproductive hormones occurred 1-2 months earlier with the peak in testicular size in September or October. In the two southern breeds (Portland and Merino), the early onset of testicular activity was more extreme with the seasonal maximum in August. In cross-bred rams, produced by mating Soay ewes (highly seasonal breed) with Portland or Merino rams (less seasonal breeds), there was a seasonal reproductive cycle that was intermediate compared to that of the parents. A comparison between all 11 breeds showed a significant correlation between the timing of the seasonal cycle in plasma FSH concentration and testicular diameter (time of peak FSH vs testis, r = 0.95). The overall results in the rams are consistent with a primary role of FSH in dictating the seasonal cycle in testicular size and the secretion of inhibin. The earlier seasonal onset in the testicular cycle in the southern breeds of domesticated sheep, and the differences from the wild type, are taken to represent the effects of genetic selection for a longer mating season.

Animals↗

Responses to selection for postmeiotic segregation frequencies in Ascobolus immersus.

A composite cross was made between 12 strains of the fungus Ascobolus immersus, six with wild-type red ascospores (w1+) and six with white ascospore mutation w1-78. A high postmeiotic segregation (PMS) frequency line was set up from colonies from ascospores from dehisced octads showing PMS, 5+ : 3w and 3+ : 5w. A low PMS line was started from ascospores from 4+ : 4w or 6+ : 2w octads, and a 'no selection' line was set up from ascospores from random octads. Colonies were crossed to tester strains to determine PMS frequencies and the selected lines were continued from ascospores of crosses of the red ascospore strain with the most extreme (e.g. high for the high line) PMS frequency with the white-ascospore strain of most extreme PMS frequency and of opposite mating type. Significant responses to selection were obtained for increased (+100%) and decreased (-58%) PMS, giving a 4.8-times difference in generation 4, with little change in the frequencies of conversion classes showing meiotic segregation (6+ : 2w and 2+ : 6w). The continuous, symmetrical, roughly normal distributions for PMS frequencies obtained when generation 5 strains were crossed to unselected tester strains are those expected if PMS frequencies are controlled by a number of polygenes, not major genes. Crosses of selected fifth-generation red-ascospore strains with extreme PMS values to base-substitution mutant w1-78, to frame-shift mutant w1-3C1 and to white-ascospore mutants w-BHj and w-9 at two loci unlinked to w1 showed that the effects of selection were not allele specific, locus specific or mutation-type specific.

Ascomycota↗

Carotenoid availability affects the development of a colour-based mate preference and the sensory bias to which it is genetically linked.

Regardless of their origins, mate preferences should, in theory, be shaped by their benefits in a mating context. Here we show that the female preference for carotenoid colouration in guppies (Poecilia reticulata) exhibits a phenotypically plastic response to carotenoid availability, confirming a key prediction of sexual selection theory. Earlier work indicated that this mate preference is genetically linked to, and may be derived from, a sensory bias that occurs in both sexes: attraction to orange objects. The original function of this sensory bias is unknown, but it may help guppies find orange-coloured fruits in the rainforest streams of Trinidad. We show that the sensory bias also exhibits a phenotypically plastic response to carotenoid availability, but only in females. The sex-specificity of this reaction norm argues against the hypothesis that it evolved in a foraging context. We infer instead that the sensory bias has been modified as a correlated effect of selection on the mate preference. These results provide a new type of support for the hypothesis that mate preferences for sexual characters evolve in response to the benefits of mate choice--the alternatives being that such preferences evolve entirely in a non-mating context or in response to the costs of mating.

Analysis of Variance↗

Evasion of mating disruption in Ephestia cautella (Walker) by increased pheromone production relative to that of undisrupted populations.

The effect of applying high dosages of pheromone for continuous disruption of mating on sex pheromone production, fecundity, and life span was studied in inbred pairs of the almond moth (Ephestia cautella) mated in glass jars. Two experiments were conducted for six and nine consecutive generations of disruption, respectively, with a pheromone mixture (2.2-2.4mg/jar in the first experiment and 10mg/jar in the second experiment) composed of two components, (Z,E)-9,12-tetradecadienyl acetate (D) and (Z)-9-tetradecenyl acetate (M), in various proportions. In the first experiment pheromone emission decreased considerably in the first generation of disruption, an average of 35% lower in the control F(1) females and 47% lower amount in females of the disrupted population (exposed to an excessive pheromone dose) as compared with the amount recorded in parental (P) females (1252+/-84pg). The reduction in pheromone emission continued in the two treatment groups, by an average of 92 pg/generation in the control and 64pg/generation under disruption until F(5). The same trend of reduction of pheromone production (as measured in gland extract) was also detected in the second experiment, with leveling off at 2.5+/-2.3 and 2.7+/-1.6ng/gland in F(7) and F(8), respectively, for the control population, and 5.7+/-5.6 and 5.9+/-5.7ng/gland in F(7) and F(8), respectively, for the disrupted population (initial pheromone level in P was 33.8+/-18.7ng/gland). The confinement of the pairs in closed jars was probably the major cause for the reduction in pheromone production besides inbreeding. The higher pheromone titer in the disrupted population relative to that in the control population may represent an evasion of the pressure of mating disruption caused by selection. The mean number of progeny (per jar) gradually decreased in successive generations in both the control and disruption treatments, but no significant difference was found between the number of progeny averaged over all the generations in the two treatments. For a pheromone blend having the average composition of D:M=77:23 in the first experiment, there was a small but significant increase in the life span among disrupted mated and unmated males (10.5 and 10.2 days, respectively) and among disrupted mated and unmated females (7.7 and 11.0 days) as compared with the respective non-disrupted control groups (9.3 and 6.5 days for the males and 6.7 and 8.6 days for the females).

Journal Article↗

Chromosome mobilization by the R plasmid R68.45: a tool in Pseudomonas genetics.

The conjugative plasmid R68.45 mobilizes the chromosome of Pseudomonas aeruginosa strain PAO from multiple sites located in different chromosome regions. In interrupted matings on the plate, selection for any single marker tested resulted in entry times of 3-5 min. When selection was imposed for two markers linked in R68.45-mediated conjugation, double recombinants appeared after a delay which corresponded approximately to the map distance between the two markers as measured by the sex factor FP2. Thus, R68.45 and FP2 appear to promote chromosome transfer at similar rates, but R68.45, unlike FP2, seems to give non-polarized transfer. R68.45 may be used to estimate map distances between linked markers located in those chromosome regions where other sex factors do not produce enough recombinants to permit accurate measurement of entry times. In R68.45 matings on the plate, most recombinants inherited short donor chromosome fragments (usually less than 10 min long) and lost the R plasmid during purification. Used like a "large" generalized transducing phage, R68.45 has proved valuable in construction of PAO strains with desired genotypes.

Chromosomes, Bacterial↗

Quantitative genetic models of female choice based on "arbitrary" male characters.

Multivariate, quantitative genetic models are developed for the evolution of female mating preferences in situations where males contribute only their gametes to their progeny. Although female mating preferences may not be directly subject to selection, they can evolve via genetic correlations with other characters that are undergoing evolutionary change. The first set of models examines the evolutionary origin of mating preferences directed at one or more traits that may or may not be expressed only in males. When several selected characters possess additive genetic variance, an indirect selective force exists for the evolution of multivariate mating preferences. The magnitude of this force is proportional to the covariance between a female's relative preference for a given male's phenotype and the expected viability of his progeny. The contribution of any single character to this covariance determines its potential value as a mate choice criterion. The pattern of genetic and phenotypic covariation may cause selectively unimportant traits to be useful in mate choice. In the extreme, selectively neutral characters may become the objects of mating preferences, if they are relatively immune to random environmental variation and genetically correlated with selectively important characters. The second set of models examines the dynamic evolution of such a selectively neutral ("arbitrary") character that is both the object of a mating preference and genetically correlated with a third trait that affects viability. The outcome of evolution in this three character system is highly indeterminate. As in other sexual selection models, there exists a line of neutral equilibria wherein the mean of the criterion character matches the mean level of mating preference within the population, while the viability trait equilibrates at the phenotypic value conferring maximum viability. This line of equilibria, however, is not likely to be stable unless females choose mates according to absolute mating preferences. Thus, mating preferences that initially may arise as a mean of increasing offspring viability may nevertheless lead to indeterminate and potentially maladaptive evolutionary outcomes.

Animals↗

Effects of population size and selection intensity of short-term response to selection for postweaning gain in mice.

The effects of population size and selection intensity on the mean response was examined after 14 generations of within full-sib family selection for postweaning gain in mice. Population sizes of 1, 2, 4, 8 and 16 pair matings were each evaluated at selection intensities of 100% (control), 50% and 25% in a replicated experiment. Selection response per generation increased as selection intensity increased. Selection response and realized heritability tended to increase with increasing population size. Replicate variability in realized heritability was large at population sizes of 1, 2 and 4 pairs. Genetic drift was implicated as the primary factor causing the reduced response and lowered repeatability at the smaller population sizes. Lines with intended effective population sizes of 62 yielded larger selection responses per unit selection differential than lines with effective population sizes of 30 or less.

Analysis of Variance↗