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Natural selection and the reinforcement of mate recognition.

Natural selection on mate recognition may often contribute to speciation, resulting in reproductive character displacement. Field populations of Drosophila serrata display reproductive character displacement in cuticular hydrocarbons when sympatric with Drosophila birchii. We exposed field sympatric and allopatric populations of D. serrata to experimental sympatry with D. birchii for nine generations. Cuticular hydrocarbons of field allopatric D. serrata populations evolved to resemble the field sympatric populations, whereas field sympatric D. serrata populations remained unchanged. Our experiment indicates that natural selection on mate recognition resulted in the field pattern of reproductive character displacement.

Analysis of Variance↗

Female life span and fertility are increased by the ejaculates of preferred males.

In animals with internal fertilization, sperm competition among males can favor the evolution of male ejaculate traits that are detrimental to females. Female mating preferences, in contrast, often favor traits in males that are beneficial to females, yet little is known about the effect of these preferences on the evolution of male ejaculates. A necessary condition for female preferences to affect the evolution of male ejaculate characteristics is that females select mates based on a trait correlated with ejaculate quality. Previous work has shown that females of the variable field cricket, Gryllus lineaticeps, prefer males that produce calling songs containing faster and longer chirps. In this study, we tested the hypothesis that females receive more beneficial ejaculates from preferred males. Females were placed on either a high- or a reduced-nutrition diet then mated twice to a male of known song phenotype. Females received only sperm and seminal fluid from males during these matings. There was no effect of male song phenotype on any fitness component for females on the high-nutrition diet. Reduced-nutrition females mated to males that produced preferred song types, however, lived longer, produced more eggs, produced more fertile eggs, and had a higher proportion of their eggs fertilized than those mated to other males. The life-span benefit was positively associated with male chirp duration, and the reproductive benefits were positively associated with male chirp rate. We explored two possible mechanisms for the life span and reproductive benefits. First, a path analysis suggested that part of the effect of male chirp duration on female life span may have been indirect; females mated to males that produced longer chirps showed delayed oviposition, and females that delayed oviposition lived longer. Males that produce longer chirps may thus transfer fewer or less potent oviposition stimulants to females in their seminal fluid. Second, there was a positive correlation between male chirp rate and the number of sperm transferred to females. The fertility benefit may thus have resulted from females receiving more sperm from males that produce faster chirps. Finally, there was a negative phenotypic correlation between male chirp rate and chirp duration, suggesting that females may have to trade off the life span and reproduction benefits when selecting a mate.

Animal Communication↗

Genetic constraints on the evolution of mate recognition under natural selection.

Field populations of Drosophila serrata display reproductive character displacement in cuticular hydrocarbons (CHCs) when sympatric with Drosophila birchii. We have previously shown that the naturally occurring pattern of reproductive character displacement can be experimentally replicated by exposing field allopatric populations of D. serrata to experimental sympatry with D. birchii. Here, we tested whether the repeated evolution of reproductive character displacement in natural and experimental populations was a consequence of genetic constraints on the evolution of CHCs. The genetic variance-covariance (G) matrices for CHCs were determined for populations of D. serrata that had evolved in either the presence or absence of D. birchii under field and experimental conditions. Natural selection on mate recognition under both field and experimental sympatric conditions increased the genetic variance in CHCs consistent with a response to selection based on rare alleles. A close association between G eigenstructure and the eigenstructure of the phenotypic divergence (D) matrix in natural and experimental populations suggested that G matrix eigenstructure may have determined the direction in which reproductive character displacement evolved during the reinforcement of mate recognition.

Animals↗

Up-regulated TGF-beta mRNA expression in splenic T cells of high IgA-prone mice: a murine model of IgA nephropathy with glomerulosclerosis.

BACKGROUND/AIMS: Recently, we established a high serum IgA-prone inbred (HIGA) mouse strain as a murine model of spontaneous IgA nephropathy by selective mating of high serum IgA ddY mice, and found that they showed enhanced production of glomerular extracellular matrix components with increased expression of TGF-beta mRNA and protein in the kidneys. In this study, we examined the roles of lymphocytes in the development of high serum IgA in this strain. METHODS: We performed flow cytometric analyses of T and B cells in splenic mononuclear cells (SMNCs) from these mice using BALB/c mice as normal controls. We also compared serum TGF-beta1 concentrations and TGF-beta mRNA expression levels in the B-cell-depleted (T-cell-rich) fraction of SMNCs in these mice. RESULTS: HIGA mice showed significantly fewer CD3-positive cells compared with BALB/c mice when young, but not when aged. The CD4/CD8 ratio of HIGA mice was lower than that of BALB/c mice, but this difference was not significant. Although the number of B220-positive cells did not vary significantly, the ratio of surface IgA-positive B cells was significantly increased in both young and adult HIGA mice. The B-cell-depleted SMNCs from HIGA mice exhibited higher levels of expression of TGF-beta and TGF-beta1 mRNA than controls from a young age, which were maintained throughout life, but there were no differences in PDGF, MCP-1 or bFGF expression between these two strains. In contrast to local mRNA expression, serum TGF-beta1 concentration was decreased in HIGA mice compared with BALB/c controls. CONCLUSION: These findings suggest that the mating procedure performed to establish HIGA mice selected for a unique phenotype of local up-regulation of TGF-beta production in the kidneys, as well as T cells that may contribute to both the early and consistently high serum IgA expression and enhanced production of renal extracellular matrix components in HIGA mice. Additionally, TGF-beta1 may act locally, not systemically, in a paracrine or autocrine manner.

Animals↗

Genetic response and inbreeding with different selection methods and mating designs for nucleus breeding programs of dairy cattle.

Stochastic simulation was used to study the effect of selection and mating strategy on rates of genetic response and inbreeding with a closed nucleus breeding program for juvenile and adult schemes with 8 males and 64 females selected to produce 1024 progeny (512 females). Selection strategies considered using all available information or only individual and sibling records. Selection of sires was either unrestricted or restricted to between full-sib families. The effect of avoidance of mating of relatives to limit inbreeding was also evaluated. Four mating designs were examined: each dam was mated to 1, 2, 4, or all sires. Mating designs involving one sire per dam and more than one dam per sire were referred to as hierarchical. Use of several mates per dam resulted in a factorial mating design. Selected parents were mated either randomly, best to best, or best to worst. An index based on relative inbreeding to response ratio was used to describe the effectiveness of strategies for reducing inbreeding relative to changes in rates of genetic response. Strategies that lower index values were preferred and include selection on BLUP or approximations of BLUP and factorial mating designs that involve the random mating of dams to several sires. Factorial mating designs were effective for a range of heritabilities. Avoidance of matings of full sibs and restriction of selection of sires to between full-sib families enabled appreciable reductions in the index. Nucleus breeding programs based entirely on the selection of juveniles were not indicated because they had higher index values than adult schemes.

Animals↗

The strength of indirect selection on female mating preferences.

An important but controversial class of hypotheses concerning the evolution of female preferences for extreme male mating displays involves "indirect selection." Even in the absence of direct fitness effects, preference for males with high overall fitness can spread via a genetic correlation that develops between preference alleles and high fitness genotypes. Here we develop a quantitative expression for the force of indirect selection that (i) applies to any female mating behavior, (ii) is relatively insensitive to the underlying genetics, and (iii) is based on measurable quantities. In conjunction with the limited data now available, it suggests that the evolutionary force generated by indirect selection on preferences is weak in absolute terms. This finding raises the possibility that direct selection on preference genes may often be more important than indirect selection, but more data on the quantities identified by our model and on direct selection are needed to decide the question.

Animals↗

Mole rat spalax ehrenbergi: mating behavior and its evolutionary significance.

Mating behavior of the subterranean mole rat, Spalax ehrenbergi, consists of three distinct stages-agonistic, courtship, and copulation. Spalax sexual behavior reflects certain cricetid affinities, some features general in rodents, and others presumably related to its subterranean, territorial life. Within four groups of Spalax ehrenbergi, each with different numbers of chromosomes, recently found in Israel, mating behavior seems to provide partial reproductive barriers. Selective matings between chromosome forms may complement a cytologic isolating mechanism to prevent widespread natural hybridization.

Animals↗

Direct and correlated responses to artificial selection on male mating frequency in the stalk-eyed fly Cyrtodiopsis dalmanni.

Traditionally it was thought that fitness-related traits such as male mating frequency, with a history of strong directional selection, should have little additive genetic variance and thus respond asymmetrically to bidirectional artificial selection. However, recent findings and theory suggest that a balance between selection for increased male mating frequency and opposing selection pressures on physiologically linked traits will cause male mating frequency to have high additive genetic variation and hence respond symmetrically to selection. We tested these hypotheses in the stalk-eyed fly, Cyrtodiopsis dalmanni, in which males hold harems comprising many females and so have the opportunity to mate at extremely high frequencies. We subjected male stalk-eyed flies to artificial selection for increased ('high') and decreased ('low') mating frequency in the presence of ecologically realistic, high numbers of females. High line males mated significantly more often than control or low line males. The direct response to selection was approximately symmetric in the high and low lines, revealing high additive genetic variation for, and no significant genetic constraints on, increased male mating frequency in C. dalmanni. In order to investigate trade-offs that might constrain male mating frequency under natural conditions we examined correlated responses to artificial selection. We measured accessory gland length, testis length and eyespan after 7 and 14 generations of selection. High line males had significantly larger accessory glands than low line males. No consistent correlated responses to selection were found in testis length or eyespan. Our results suggest that costs associated with the production and maintenance of large accessory glands, although yet to be identified, are likely to be a major constraint on mating frequency in natural populations of C. dalmanni.

Animals↗

Spouse resemblance in body mass index: effects on adult obesity prevalence in the offspring generation.

Accruing evidence indicates that mate selection is promoted by similarity in body fatness. Assortative mating for obesity may contribute genetically to the obesity epidemic by increasing the risk in subsequent generations. To test this hypothesis, the authors analyzed measured and validated questionnaire data on family members, obtained between 1987 and 2000 from 7,834 obese probands and from 829 subjects randomly ascertained from the general Swedish population. Spouse correlations in body mass index were strongest among couples with the shortest duration of cohabitation. Obesity concordance in parents was associated with an obesity prevalence of 20.1% in adult offspring compared with 1.4% if parents were concordantly nonobese (odds ratio = 18.3, 95% confidence interval: 9.0, 37.4). The prevalence was 8.2% if parents were obesity discordant (odds ratio = 6.5, 95% confidence interval: 3.2, 13.2). No association was found between rearing parents' and nonbiologic offspring's body mass index. These results agree with the hypothesis that assortative mating for obesity confers a higher risk of obesity in the offspring generation and thus contributes to the obesity epidemic. Parental obesity concordance is a strong, easily identifiable genetic risk factor that should be considered in the complex network of risk factors for obesity in designing primary prevention programs.

Adult↗

Reproductive character displacement of male stickleback mate preference: reinforcement or direct selection?

Reproductive isolation can evolve between species as a byproduct of adaptation to different niches, through reinforcement, and by direct selection on mating preferences. We investigated the potential role of direct selection in the reproductive isolation between sympatric species of threespine sticklebacks. Each sympatric pair consists of a small-bodied limnetic species and large-bodied benthic species. We compared the mate preferences and courtship behavior of males from one sympatric limnetic population and two allopatric populations. We used limnetic-like allopatric populations to control for the effects of ecological character displacement and adaptation to different niches on mate preferences. The sympatric limnetic males preferred the small limnetic females, whereas the allopatric limnetic-like males preferred the large benthic females, suggesting that adaptation to the limnetic niche does not automatically confer a preference for small limnetic females. This reproductive character displacement of male preference is consistent with the predictions of both reinforcement and direct selection on mate preferences. To test for direct selection, we assessed a prediction of one proposed mechanism: predation by benthic females on eggs guarded by limnetic males. The allopatric males come from populations in which there is no egg predation. Sympatric limnetic males were more aggressive toward benthic females than toward limnetic females, whereas the allopatric limnetic-like males did not treat the two types of females differently. The contrast in male behavior suggests that egg predation has shaped male preferences. Direct selection is potentially more effective than indirect selection via reinforcement, and it is likely that it has been important in building up reproductive isolation between limnetic and benthic sticklebacks.

Adaptation, Physiological↗

Conjugational recombination in Escherichia coli: genetic analysis of recombinant formation in Hfr x F- crosses.

The formation of recombinants during conjugation between Hfr and F- strains of Escherichia coli was investigated using unselected markers to monitor integration of Hfr DNA into the circular recipient chromosome. In crosses selecting a marker located approximately 500 kb from the Hfr origin, 60-70% of the recombinants appeared to inherit the Hfr DNA in a single segment, with the proximal exchange located > 300 kb from the selected marker. The proportion of recombinants showing multiple exchanges increased in matings selecting more distal markers located 700-2200 kb from the origin, but they were always in the minority. This effect was associated with decreased linkage of unselected proximal markers. Mutation of recB, or recD plus recJ, in the recipient reduced the efficiency of recombination and shifted the location of the proximal exchange(s) closer to the selected marker. Mutation of recF, recO or recQ produced recombinants in which this exchange tended to be closer to the origin, though the effect observed was rather small. Up to 25% of recombinant colonies in rec+ crosses showed segregation of both donor and recipient alleles at a proximal unselected locus. Their frequency varied with the distance between the selected and unselected markers and was also related directly to the efficiency of recombination. Mutation of recD increased their number by twofold in certain crosses to a value of 19%, a feature associated with an increase in the survival of linear DNA in the absence of RecBCD exonuclease. Mutation of recN reduced sectored recombinants in these crosses to approximately 1% in all the strains examined, including recD. A model for conjugational recombination is proposed in which recombinant chromosomes are formed initially by two exchanges that integrate a single piece of duplex Hfr DNA into the recipient chromosome. Additional pairs of exchanges involving the excised recipient DNA, RecBCD enzyme and RecN protein, can subsequently modify the initial product to generate the spectrum of recombinants normally observed.

Chromosome Mapping↗

Female mating behaviour, sexual selection and chromosome I inversion karyotype in the seaweed fly, coelopa frigida

Previous studies of the seaweed fly, Coelopa frigida, have revealed the operation of several different forces of sexual selection. The overall pattern of mate choice seen in natural populations is not consistent with the predictions of indirect sexual selection as females do not express preferences that maximize the fitness of their offspring, even though the benefits from such choice are relatively large in this species. Thus, the maintenance of female mate choice for large male size must instead either be a result of a direct benefit to the female of mating with large males, or a side-effect of the evolution of another character, in other words pleiotropy. In order to separate these two alternatives the genetical basis of female mating behaviour needs to be studied. Previous studies have revealed associations between chromosomal inversion karyotype and both general female willingness to mate and mate choice for large male size, however these associations were lost after several generations of laboratory culture. Here several isokaryotypic lines from wild collections of flies were derived. The willingness to mate and mate choice of females from each line were determined. Pairs of lines of opposite inversion karyotype that significantly differed in either or both willingness to mate and mate choice were crossed. The mating behaviour and inversion karyotype of the F2 progeny (all F1 progeny are heterokaryotypes) were determined. Clear differences in the general levels of female willingness to mate were found between the two inversion homokaryotypes in several families, whereas variation in the strength of female choice for large male size were not revealed. It is suggested that mate choice in this species occurs as a pleiotropic effect of selection acting on female willingness to mate.

Journal Article↗

Evolutionary pathways in shorebird breeding systems: sexual conflict, parental care, and chick development.

Sexual selection, mating opportunities, and parental behavior are interrelated, although the specific nature of these relationships is controversial. Two major hypotheses have been suggested. The parental investment hypothesis states that the relative parental investment of the sexes drives the operation of sexual selection. Thus, the sex that invests less in offspring care competes more intensely and monopolizes access to mates. The sexual conflict hypothesis proposes that sexual selection (the competition among both males and females for mates), mating opportunities, and parental behavior are interrelated and predicts a feedback loop between mating systems and parental care. Here we test both hypotheses using a comprehensive dataset of shorebirds, a maximum-likelihood statistical technique, and a recent supertree of extant shorebirds and allies. Shorebirds are an excellent group for these analyses because they display unique variation in parental care and social mating system. First, we show that chick development constrains the evolution of both parental care and mate competition, because transitions toward more precocial offspring preceded transitions toward reduced parental care and social polygamy. Second, changes in care and mating systems respond to one another, most likely because both influenced and are influenced by mating opportunities. Taken together, our results are more consistent with the sexual conflict hypothesis than the parental investment hypothesis.

Animals↗

Ontogeny of aggressive-sexual behavior between males in lines of Japanese quail (Coturnix coturnix japonica) selected for male mating frequency.

Japanese quail from lines bidirectionally selected for high and low male mating frequency and from the random-bred base population were observed in an ontogenetic study of aggressive-sexual behavior. Quail were reared in sex-intermingled flocks until 28 days of age at which time half of the males from each line were housed as all-male flocks and half were transferred to individual cages. At 41 days of age, individually caged males exhibited mounting behavior to win encounters with other males. By 45 days, males from the high and control lines exhibited significantly more aggressive-sexual behavior than those from the low lines. Line x Rearing Experience interactions for mounting behavior at 45 and 56 days of age were due to the individually caged high- and control-line males' winning encounters by mounts; those maintained in flocks rarely mounted. Although some low-line males won encounters by mounts, occurrence of this behavior was infrequent. Males maintained in flocks were placed in individual cages at 57 days of age. When these males were compared with those housed in individual cages from 28 days of age, at 84 days of age and older, the Line x Rearing Experience interaction observed previously disappeared, and only differences among genetic lines were evident. Genetic and rearing experience effects are discussed as influencing agonistic and sexual behaviors.

Age Factors↗

The importance of calling song and courtship song in female mate choice in the variable field cricket.

Male field crickets produce calling songs, courtship songs, tactile signals and chemical signals. Although calling songs are known to play an important role in female mate choice, the importance of the other signals in mate choice is poorly understood. In the variable field cricket, Gryllus lineaticeps, females select mates, in part, based on variation in male calling song. Females prefer higher chirp rates, a trait which is partially dependent on male nutrient intake, and females prefer longer chirp durations, a trait which appears to be independent of male nutrient intake. We tested whether females also have preferences based on variation in male courtship song, and whether the structure of male courtship song varies with nutrient intake. First, we reexamined female preference for calling song chirp rate. Then, we examined: (1) female preference based on courtship song chirp rate; (2) the relative importance of calling song and courtship song chirp rate; (3) the nutrition dependence of courtship song chirp rate; and (4) the correlation between calling song and courtship song chirp rate. As reported previously, females preferred higher calling song chirp rates, and in addition, preferred higher courtship song chirp rates. Females were more likely to switch from a speaker broadcasting more attractive calling song to a speaker broadcasting less attractive calling song when the attractive calling song was associated with an unattractive courtship song than when it was associated with an attractive courtship song. Preferences based on courtship song may thus cause females to alter the choices that they made based on calling song. Males that received greater nutrients did not produce higher courtship song chirp rates. There was no correlation between calling song and courtship song chirp rate. As a result, the two traits may provide information to females about different aspects of male quality. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

The shape of female mating preferences.

The "shape" of a female mating preference is the relationship between a male trait and the probability of acceptance as a mating partner. The shape of preferences is important in many models of sexual selection, mate recognition, communication, and speciation, yet it has rarely been measured precisely. Here I examine preference shape for male calling song in a bushcricket (katydid). Preferences change dramatically between races of a species, from strongly directional to broadly stabilizing (but with a net directional effect). Preference shape generally matches the distribution of the male trait. This is compatible with a coevolutionary model of signal-preference evolution, although it does not rule out an alternative model, sensory exploitation. Preference shapes are shown to be genetic in origin.

Animals↗

Good vs complementary genes for parasite resistance and the evolution of mate choice.

BACKGROUND: Female mate choice may be adaptive when males exhibit heritable genetic variation at loci encoding resistance to infectious disease. The Hamilton-Zuk hypothesis predicts that females should assess the genetic quality of males by monitoring traits that indicate health and vigor (condition-dependent choice, or CD). Alternatively, some females may employ a more direct method of screening and select mates based on the dissimilarity of alleles at the major histocompatibility loci (we refer to this as opposites-attract, or OA). Empirical studies suggest that both forms of mate choice exist, but little is known about the potential for natural selection to shape the two strategies in nature. RESULTS: We used computer simulation models to examine the evolutionary fates of the two forms of mate choice in populations at risk for infection by debilitating parasites. We found that populations exhibiting random mating (no mate choice) can be invaded and replaced completely by individuals practicing CD type mate choice. We also found that an allele encoding OA choice can increase when rare in randomly mating populations, but that it does not go to fixation under selection. A similar result was obtained when the OA strategy was introduced into populations practicing CD mate choice. As before, we found that the OA choice allele will increase when rare, and that it will not go to fixation under selection. The converse however was not true, as CD individuals gain no rare advantage when introduced into an OA population. CONCLUSIONS: Taken together, the results suggest that, when rare, OA is the best strategy for parasite evasion (of those considered here). The consequence of OA increasing in the population, however, is to reduce the parasite driven genotype oscillations and facilitate the breakdown of linkage disequilibrium at the disease-resistance loci. This leads to a neutrally stable situation in which different strategies have equal fitness, and suggests that multiple forms of mate choice may be expected to occur in populations at risk from infectious disease.

Alleles↗

Mate choice on fallow deer leks.

Leks, on which males defend small clustered mating territories, may have evolved because of the unusual opportunities they provide for female choice of mating partners, and several studies of lek-breeding animals have demonstrated correlations between the mating success of males and their phenotype or behaviour. However, these could arise because (1) females select mates on the basis of male phenotypic traits; (2) males interfere with each other's mating attempts; or (3) females show preferences for particular mating territories, and larger or stronger males are more likely to win access to these territories. Here we report that when fallow bucks on a traditional lek were experimentally induced to change their territories, differences in the mating success of bucks persisted, whereas differences in the position of their territories relative to the centre of the lek did not. The observation that bucks rarely interfered with their neighbours' harems and females moved freely between bucks suggests that females choose their mates on the basis of male phenotype rather than territory type or location. In this population, the immediate factor affecting the movements of females between males was the size of a buck's harem.

Animals↗