An analytical model for a short-term advantage for sex

An analytical treatment is given for a model of Maynard Smith in which a short-term advantage for sex and recombination is provided by the mechanism of sib-competition. Suppose the next generation is formed by the winners of a large number of contests. Suppose a number of parents each contribute M offspring to a given contest, but the offspring of an asexual parent are identical whereas those of a sexual parent are distributed with some variance. If M is large there is a high probability that a sexual offspring will have a high enough fitness to win the contest. Calculations show that values of M around 3 and 4 are generally enough for sexual behaviour to overcome its two-fold disadvantage.     

A short-term advantage for sex and recombination through sib-competition

A model in which sexual reproduction confers an immediate selective advantage is presented. The essential assumptions of the model are that the environment consists of patches with unpredictable selective features, that selection within a patch is density dependent, and that several off-spring of a single female compete within a single patch. Computer simulation shows that sexual reproduction can have a twofold advantage per generation over parthenogenesis, but this requires a total intensity of selection, $\text{W}{}_{\mathrm{<mtext>max</mtext>}}\text{?}\text{W}$ of the order of 30 or more. Alleles for higher recombination increase in frequency within a sexual population. Selection in favour of sex or of recombination is reduced or reversed if different selective features of a patch are correlated, or if several gene loci are concerned with adaptation to a single selective feature. The results can be understood qualitatively once it is appreciated that density-dependent competition between siblings is wasteful from the point of view of the parent, and that this competition becomes more intense as siblings become genetically more similar.     

Rare male advantage and sex ratio

Relative success of mutant white and wild-type CS Drosophila melanogaster males is frequency-dependent, if sex ratio is 1:1. If the number of females is constant, this success depends on the ratio between the mutant and wild-type males. The sex ratio changes strongly affect the male mating activity of both genotypes.Emphasis is placed on the general interest of the Ayala equation and the Wattiaux-Lichtenberger equation in frequency-dependence and sex ratio dependence analysis.     

Mutation-selection balance and the evolutionary advantage of sex and recombination

Mutation-selection balance in a multi-locus system is investigated theoretically, using a modification of Bulmer's infinitesimal model of selection on a normally-distributed quantitative character, taking the number of mutations per individual (n) to represent the character value. The logarithm of the fitness of an individual with n mutations is assumed to be a quadratic, decreasing function of n. The equilibrium properties of infinitely large asexual populations, random-mating populations lacking genetic recombination, and random-mating populations with arbitrary recombination frequencies are investigated. With 'synergistic' epistasis on the scale of log fitness, such that log fitness declines more steeply as n increases, it is shown that equilibrium mean fitness is least for asexual populations. In sexual populations, mean fitness increases with the number of chromosomes and with the map length per chromosome. With 'diminishing returns' epistasis, such that log fitness declines less steeply as n increases, mean fitness behaves in the opposite way. Selection on asexual variants and genes affecting the rate of genetic recombination in random-mating populations was also studied. With synergistic epistasis, zero recombination always appears to be disfavoured, but free recombination is disfavoured when the mutation rate per genome is sufficiently small, leading to evolutionary stability of maps of intermediate length. With synergistic epistasis, an asexual mutant is unlikely to invade a sexual population if the mutation rate per diploid genome greatly exceeds unity. Recombination is selectively disadvantageous when there is diminishing returns epistasis. These results are compared with the results of previous theoretical studies of this problem, and with experimental data.     

Size advantage may not always favor sex change

A simple application of sex allocation theory to sex reversal suggests that, under Ghiselin's size advantage model, this form of sexuality ought to be common; actually it is quite rare. This note suggests that a sex specific size advantage may not favor sex change if the advantage is offset by other life-history tradeoffs. A few possibilities are discussed.     

Hitch-hiking parasite: a dark horse may be the real rider

Many parasites engaged in complex life cycles manipulate their hosts in a way that facilitates transmission between hosts. Recently, a new category of parasites (hitch-hikers) has been identified that seem to exploit the manipulating effort of other parasites with similar life cycle by preferentially infecting hosts already manipulated. Thomas et al. (Evolution 51 (1997) 1316) showed that the digenean trematodes Microphallus papillorobustus (the manipulator) and Maritrema subdolum (the hitch-hiker) were positively associated in field samples of gammarid amphipods (the intermediate host), and that the behaviour of Maritrema subdolum rendered it more likely to infect manipulated amphipods than those uninfected by M. papillorobustus. Here I provide experimental evidence demonstrating that M. subdolum is unlikely to be a hitch-hiker in the mentioned system, whereas the lucky candidate rather is the closely related but little known species, Microphallidae sp. no. 15 (Parassitologia 22 (1980) 1). As opposed to the latter species, Maritrema subdolum does not express the appropriate cercarial behaviour for hitch-hiking.     

No evidence of short-term exchange of meat for sex among chimpanzees

The meat-for-sex hypothesis posits that male chimpanzees (Pan troglodytes) trade meat with estrous females in exchange for short-term mating access. This notion is widely cited in the anthropological literature and has been used to construct scenarios about human evolution. Here we review the theoretical and empirical basis for the meat-for-sex hypothesis. We argue that chimpanzee behavioral ecology does not favor the evolution of such exchanges because 1) female chimpanzees show low mate selectivity and require little or no material incentive to mate, violating existing models of commodity exchange; and 2) meat-for-sex exchanges are unlikely to provide reproductive benefits to either partner. We also present new analyses of 28 years of data from two East African chimpanzee study sites (Gombe National Park, Tanzania; Kanyawara, Kibale National Park, Uganda) and discuss the results of previously published studies. In at least three chimpanzee communities, 1) the presence of sexually receptive females did not increase hunting probability, 2) males did not share preferentially with sexually receptive females, and 3) sharing with females did not increase a male's short-term mating success. We acknowledge that systematic meat sharing by male chimpanzees in expectation of, or in return for, immediate copulations might be discovered in future studies. However, current data indicate that such exchanges are so rare, and so different in nature from exchanges among humans, that with respect to chimpanzees, sexual bartering in humans should be regarded as a derived trait with no known antecedents in the behavior of wild chimpanzees.     

Emerging sex allocation research in mammals: marsupials and the pouch advantage

• 1 Adaptive adjustments in offspring sex ratios in mammals have long been reported, but the conditions and mechanisms that prompt shifts in the proportion of sons and daughters born are still unclear.
• 2 Empirical evidence indicates that offspring sex in mammals can be related to a diversity of environmental and maternal traits. However, the underlying assumptions regarding offspring and maternal fitness are rarely tested.
• 3 Physiological mechanisms of maternal selection of offspring sex may occur at many stages during the prolonged maternal investment stage, and a pluralistic approach to studying mechanisms might prove fruitful.
• 4 This review highlights the apparent frequency, in marsupial mammals, of sex ratio bias, which has largely been recorded as conforming to one of a few hypotheses.
• 5 Marsupials are ideally suited to experiments involving cross‐fostering of offspring, which can allow rigorous tests of the fitness consequences of rearing one sex vs. the other. The reproductive biology of marsupials lends the group to detailed studies of the timing and physiological correlates of offspring sex biases.
• 6 Many components of metatherian biology may prove advantageous in experimental studies of sex allocation in mammals, and together may provide a prosperous avenue for examining adaptive and mechanistic hypotheses in mammalian sex allocation.

     

Inbreeding,heterozygote advantage and the evolution of neo-X and neo-Y sex chromosomes

Associations between heterozygosities at different loci are generated by inbreeding. This can cause a fusion or translocation involving a sex chromosome and an autosome to have a selective advantage, when there is selection in favour of heterozygotes. Population genetic models of Y-autosome and X-autosome rearrangments in populations mating by a mixture of full sib-matings and random mating are described, in which the rearrangements cause an autosomal locus with heterozygote advantage to become linked to the true sex chromosomes. Such rearrangements gain a selective advantage under a wide range of conditions. If they can invade, Y-autosome rearrangements always spread to fixation, whereas X-autosome rearrangements may be maintained as stable polymorphisms. The results are discussed in relation to data on breeding systems and karyotypic evolution in termites.     

Drift increases the advantage of sex in RNA bacteriophage Phi6

The pervasiveness of sex and recombination remains one of the most enigmatic problems in evolutionary biology. According to many theoretical models, recombination can increase the rate of adaptation by restoring genetic variation. However, the potential for genetic drift to generate conditions that produce this outcome has yet to be studied experimentally. We have designed and performed an experiment that reveals the effects of drift on existing genetic variation by minimizing the influence of variation on beneficial mutation rate. Our experiment was conducted in populations of RNA bacteriophage Phi6 initiated from a common source population at varying bottleneck sizes. The segmented genome of this virus results in genetic exchange between viruses that co-infect the same host cell. In response to selection for growth in a high-temperature environment, sexual lines outperformed their asexual counterparts on average. The advantage of sex attenuated with increasing effective population size, implying that the rate of adaptation was limited by clonal interference among segments caused by drift. This is the first empirical evidence that the advantage of sex during adaptation increases with the intensity of drift.     

Hitch-hiking: a parallel heuristic search strategy, applied to the phylogeny problem.

The article introduces a parallel heuristic search strategy ("Hitch-hiking") which can be used in conjunction with other random-walk heuristic search strategies. It is applied to an artificial phylogeny problem, in which character sequences are evolved using pseudo-random numbers from a hypothetical ancestral sequence. The objective function to be minimized is the minimum number of character-state changes required on a binary tree that could account for the sequences observed at the tips (leaves) of the tree -- the Maximum Parsimony criterion. The Hitch-hiking strategy is shown to be useful in that it is robust and that on average the solutions found using the strategy are better than those found without. Also the strategy can dynamically provide information on the characteristics of the landscape of the problem. I argue that Hitch-hiking as a scheme for parallelization of existing heuristic search strategies is of potentially very general use, in many areas of combinatorial optimization.     

Size advantage for male function and size‐dependent sex allocation in Ambrosia artemisiifolia,a wind‐pollinated plant

In wind‐pollinated plants, male‐biased sex allocation is often positively associated with plant size and height. However, effects of size (biomass or reproductive investment) and height were not separated in most previous studies. Here, using experimental populations of monoecious plants, Ambrosia altemisiifolia, we examined (1) how male and female reproductive investments (MRI and FRI) change with biomass and height, (2) how MRI and height affect male reproductive success (MRS) and pollen dispersal, and (3) how height affects seed production. Pollen dispersal kernel and selection gradients on MRS were estimated by 2,102 seeds using six microsatellite markers. First, MRI increased with height, but FRI did not, suggesting that sex allocation is more male‐biased with increasing plant height. On the other hand, both MRI and FRI increased with biomass but often more greatly for FRI, and consequently, sex allocation was often female‐biased with biomass. Second, MRS increased with both height and MRI, the latter having the same or larger effect on MRS. Estimated pollen dispersal kernel was fat‐tailed, with the maximum distance between mates tending to increase with MRI but not with height. Third, the number of seeds did not increase with height. Those findings showed that the male‐biased sex allocation in taller plants of A. artemisiifolia is explained by a direct effect of height on MRS.     

Evolution of genetic variability and the advantage of sex and recombination in changing environments.

The role of recombination and sexual reproduction in enhancing adaptation and population persistence in temporally varying environments is investigated on the basis of a quantitative-genetic multilocus model. Populations are finite, subject to density-dependent regulation with a finite growth rate, diploid, and either asexual or randomly mating and sexual with or without recombination. A quantitative trait is determined by a finite number of loci at which mutation generates genetic variability. The trait is under stabilizing selection with an optimum that either changes at a constant rate in one direction, exhibits periodic cycling, or fluctuates randomly. It is shown by Monte Carlo simulations that if the directional-selection component prevails, then freely recombining populations gain a substantial evolutionary advantage over nonrecombining and asexual populations that goes far beyond that recognized in previous studies. The reason is that in such populations, the genetic variance can increase substantially and thus enhance the rate of adaptation. In nonrecombining and asexual populations, no or much less increase of variance occurs. It is explored by simulation and mathematical analysis when, why, and by how much genetic variance increases in response to environmental change. In particular, it is elucidated how this change in genetic variance depends on the reproductive system, the population size, and the selective regime, and what the consequences for population persistence are.     

Effects of short-term administration of sex hormones on normal and autoimmune mice

The effects of short-term administration (2 to 4 wk) of sex hormones on the immune system of normal (C57BL/6) and autoimmune (C57BL/6-lpr, C3H/lpr, B/W) strains of mice were investigated. Both estrogen (E2) and testosterone (Te) had significant effects on the numbers of T and B cells as well as on the density of cell surface antigens as demonstrated by flow cytometry. For example, Te depleted Thy-1.2+ thymocytes in normal mice and brought about a shift to lower density cells. Lyt-2+ cells appeared to be the main target cells of hormonal modulation in normal and autoimmune mice. Both sex hormones significantly depleted these cells in the thymus but had differential effects in the peripheral lymphoid organs, particularly in the spleen. In general, E2 depleted Lyt-2+ cells, whereas Te increased or maintained this subpopulation of cells in spleen and lymph nodes. Similarly, the suppressor cell activity and IL 2 production on a per cell basis in E2-treated animals was diminished, whereas Te-treated animals had normal or enhanced activity. The relevance of these findings to differential sex susceptibility in autoimmune diseases is discussed.     

Classic predictions about sex change do not hold under all types of size advantage

Theory predicts that the ‘size advantage’ (rate of increase in male and female fitness with age or size) determines the direction and the timing of sex change in sequential hermaphrodites. Whereas the size advantage is generated by the mating system and would be expected to vary within and between species, the shape or form of the size advantage has rarely been estimated directly. Here, we ask whether theoretical predictions about the timing of sex change hold under different types of size advantage. We model two biological scenarios representing different processes generating the size advantage and find that different types of size advantage can produce patterns that qualitatively differ from classic predictions. Our results demonstrate that a good understanding of sequentially hermaphroditic mating systems, and specifically, a direct assessment of the processes underlying the size advantage is crucial to reliably predict and explain within‐species patterns of the timing of sex change.