Learning rules for optimal selection in a varying environment: mate choice revisited

The quality of a chosen partner can be one of the most significantfactors affecting an animal's long-term reproductive success.We investigate optimal mate choice rules in an environment wherethere is both local variation in the quality of potential mateswithin each local mating pool and spatial (or temporal) variationin the average quality of the pools themselves. In such a situation,a robust rule that works well across a variety of environmentswill confer a significant reproductive advantage. We formulatea full Bayesian model for updating information in such a varyingenvironment and derive the form of the rule that maximizes expectedreward in a spatially varying environment. We compare the theoreticalperformance of our optimal learning rule against both fixedthreshold rules and simpler near-optimal learning rules andshow that learning is most advantageous when both the localand environmental variances are large. We consider how optimalsimple learning rules might evolve and compare their evolutionwith that of fixed threshold rules using genetic algorithmsas minimal models of the relevant genetics. Our analysis pointsup the variety of ways in which a near-optimal rule can be expressed.Finally, we describe how our results extend to the case of temporallyvarying environments.     

Efficient computation of subset selection probabilities with application to Cox regression

An efficient method is presented to compute the probabilityof selection of a specified subset from the set of all subsetsof a fixed size where the subsets are taken from a populationwhose units have varying individual probabilities of selection.The problem is motivated by the computation of the exact marginallikelihood for the Cox proportional hazards model.     

Multiple cues in mate selection: The sexual interference hypothesis

Animals use multiple cues when choosing mates, but it is not yet clear why a single signal would not suffice. In this paper, drawing support from predation and “noise” effects on mate choice, marketing economics, and multiple signals models, a new hypothesis explaining multiple sexual signals is proposed: the sexual interference hypothesis. The hypothesis is based on three well-supported premises: (1) selectivity decreases when mate assessment costs increase, (2) assessment costs increase when the propagation or reception of sexual signals is more difficult, and (3) males not only exploit such circumstances by courting females when choice is more difficult, but actively interfere with females' preferences by making choice more difficult. The hypothesis argues that additional sexual signals evolve as a way for males to hinder female mate choice by interfering with the propagation and reception of other males' sexual signals. Females respond by evolving the ability to glean meaningful information from signals despite males' attempts at obfuscation. In turn, males respond by producing better interference signals and signals that are not so easily blocked. This co-evolutionary process increases the costs of assessment for females and the costs of signal production for males, and leads to a temporary equilibrium of honest advertising via multiple signals.     

Rules for assortative mating in relation to selection for linear merit functions

Summary This study examined how assortative mating (without selection) based on linear combinations of two traits could be used to change genetic parameters so as to increase efficiency of selection. The efficiency of the Smith-Hazel index for improvement of multiple traits is a function of phenotypic and genetic variances and covariances, and of the relative economic values of the traits involved. Assortative mating is known to change genetic variances and covariances. Recursive formulae were derived to obtain these variances and covariances after t generations of assortative mating on linear combinations (mating rules) of phenotypic values for two traits, with a given correlation between mates. Selection efficiency after t generations of assortative mating without selection was expressed as a function of random mating genetic parameters, economic values, the mating rule, and the correlation between mates. Selection efficiency was maximized with respect to the coefficients in the mating rule. Because the objective function was nonlinear, a computer routine was used for maximizing it. Two cases were considered. When random mating heritabilities for the two traits were h _X ² =0.25 and h _Y ² =0.50, the genetic correlation r_XY=-0.60, and the economic values were a_X=3 and a_Y=1, continued assortative mating based on the optimal mating rule for 31 generations (with a correlation of 0.80 between mates) increased selection efficiency by 29%. Heritabilities changed to 0.38 and 0.66, respectively, and the genetic correlation became – 0.79. When h _X ² =0.60, h _Y ² =0.60, r_XY=– 0.20, a₁=1 and a₂=1, 36 generations of continued assortative mating with the optimal mating rule increased the efficiency of selection by 17%, heritabilities became h _X ² = h _Y ² =0.71, and the genetic correlation changed to 0.25. Only three generations of assortative mating were required to change the sign of the genetic correlation.     

HLA and mate selection: no evidence in South Amerindians.

There have been reports of nonrandom mating (negative-assortative mating) or preference for individuals of different major histocompatibility complex (MHC) genotypes in both mice and humans. We have examined the association of HLA-A and HLA-B genotypes, both for each locus by itself and for two-locus genotypes, in mates of 194 couples from 11 South Amerindian tribes. The proportion of couples sampled averaged >50% of the total matings with progeny for 10 of the tribes. In nearly all cases, HLA-sharing proportions were very close to those expected from random mating, suggesting strong negative-assortative mating for MHC is not present in these South Amerindians.     

The evolutionary psychology of mate selection in Morocco

Patterns of mate preference in Morocco are investigated in order to test whether they support hypotheses advanced by David Buss and other evolutionary psychologists. Because of the custom of cousin marriage in Morocco, a multivariate model that included cosocialization data was developed for the purpose of testing the Westermarck hypothesis of inbreeding avoidance. Hence, two previously separate domains of research are unified in one design that permits the further exploration of questions pertaining to the domain specificity of psychological mechanisms. Multiple independent mate choice predictors were identified using logistic regression analysis. Results support the Westermarck hypothesis of inbreeding avoidance. Sleeping in the same room during childhood was found in both sexes to produce an aversion to marriage. Other evidence suggests that aversion to inbreeding extends further among females than males in that females but not males show an aversion to marriage to related individuals with whom they had daily social contact in early childhood. The evolutionary prediction that females differ from males concerning resource holding capacity was also supported. Females showed a preference for males whom they judged to have higher social status than theirs, while this criterion was unimportant for males. The predicted sex difference in preferred age of marriage partner was also supported. Contrary to previous findings, the predicted difference between the sexes with regard to physical attractiveness was not supported. This research was supported by NSF grant BSN 90-19876. Alex Walter obtained a Ph.D. in anthropology from Rutgers University in 1992. He was a research associate at Rutgers from 1994 to 1995. He has taught at Cornell College and at Rutgers and Princeton universities. He is currently working on additional articles that will present data from his dissertation research on mate selection and inbreeding avoidance in Morocco.     

Potential genetic benefits of mate selection in whitefish

To test the potential of optimal mate selection with respect to offspring viability, we crossed 10 female and 10 male whitefish in all possible combinations and reared the resulting 100 sib groups in several replicates. We recorded two types of egg mortality, one that was correlated to developmental problems, and a later one that was correlated to a bacterial infection. We found strong maternal and paternal effects in both types of mortality. Early mortality also depended on which female was mated with which male, suggesting partial incompatibilities. The later mortality, but not the former, could be predicted by male breeding ornamentation. More strongly ornamented males sired offspring that better survived the epidemic during egg development. This ‘good genes’ effect was larger than expected from theory: optimal mate selection would have improved offspring survival during the epidemic by about 12% (or reduce mortality by about 66%) as compared with random mating.     

Sexual selection and mate choice in evolutionary psychology

The importance of mate choice and sexual selection has been emphasized by the majority of evolutionary psychologists. This paper assesses three cases of work on mate choice and sexual selection in evolutionary psychology: David Buss on cross-cultural human mate preferences, Randy Thornhill and Steve Gangestad on the link between mate preferences and fluctuating asymmetry, and Geoffrey Miller on the role of Fisher’s runaway process in human evolution. A mixture of conceptual and empirical problems in each case highlights the general weakness of work in evolutionary psychology on these issues.

Sex differences in mate selection strategies: Content analyses and responses to personal advertisements in Brazil

We studied mate selection strategies as revealed in heterosexual personal advertisements published in a Brazilian newspaper, analyzing both the ads' content, with respect to the attributes that men and women offered and sought, and predictors of the number of responses that each ad received. Demands made for a prospective partner changed as a function of the age of the advertiser in predictable sex-differentiated ways: women became less demanding as they aged, whereas men became more demanding. The number of responses received by men and women as a function of age followed a similarly sex-differentiated pattern, with older women receiving fewer responses than younger women, and older men receiving more responses than younger men. In general, results of the present study provided support for our predictions. People who used personal advertisements, a relatively recent unconventional way for selecting mates, expressed conventional preferences, suggestive of evolved psychological mechanisms.     

Genetic mating system and mate selection in smallmouth bass

Mating systems are an important factor influencing the variance in reproductive success among individuals within natural populations and thus have important ecological and evolutionary implications. We used molecular pedigree reconstruction techniques with microsatellite DNA data to characterize the genetic mating system and mate selection in adult smallmouth bass spawning in Lake Opeongo. The genetic mating system of smallmouth bass in this system can be characterized as predominantly monogamous with a low rate of polygynandry particularly among larger individuals. Iteroparous individuals showed a complete absence of interannual mate fidelity, presumably due to the low annual return rate of spawning adults. Within a season, individuals from both sexes pursued additional mating opportunities with males showing greater variance in mate number than females. Female mate selection appeared to be largely random with little evidence for elevated levels of inbreeding in this population. Multiple mating females pursued additional males to whom they were less related than the first male with which they spawned within a given season, however, this pattern varied among years. The mating pattern observed in this population would likely limit the strength of sexual selection and thus could account for the lack of sexual dimorphism and the absence of alternative reproductive tactics in this species.     

Natural selection and divergence in mate preference during speciation

Sexual isolation can evolve due to natural selection against hybrids (reinforcement). However, many different forms of hybrid dysfunction, and selective processes that do not involve hybrids, can contribute to the evolution of sexual isolation. Here we review how different selective processes affect the evolution of sexual isolation, describe approaches for distinguishing among them, and assess how they contribute to variation in sexual isolation among populations of Timema cristinae stick-insects. Pairs of allopatric populations of T. cristinae living on different host-plant species exhibit greater sexual isolation than those on the same host, indicating that some sexual isolation has evolved due to host adaptation. Sexual isolation is strongest in regions where populations on different hosts are in geographic contact, a pattern of reproductive character displacement that is indicative of reinforcement. Ecological costs to hybridization do occur but traits under ecological selection (predation) do not co-vary strongly with the probability of between-population mating such that selection on ecological traits is not predicted to produce a strong correlated evolutionary response in mate preference. Moreover, F1 hybrid egg inviability is lacking and the factors contributing to reproductive character displacement require further study. Finally, we show that sexual isolation involves, at least in part, olfactory communication. Our results illustrate how understanding of the evolution of sexual isolation can be enhanced by isolating the roles of diverse ecological and evolutionary processes.     

The use of constrained selection indexes in breeding for economic merit

Summary Various methods exist for the derivation of restricted and/or desired gains selection indexes, and their use in applied breeding has been advocated. It is shown that there exists a set of implied linear economic weights for all constrained indexes and their derivation is given. Where economic weights are linear and known, a standard selection index is, by definition, optimal and thus a constrained index will usually be suboptimal. It is argued that economic weights can always be estimated and that the effects of uncertain weights can be examined by sensitivity analysis. If economic weights are nonlinear, use of the first order (linear) economic weights or a derived linear index, using previously described methods, will give very close to optimum economic selection responses. Examples from the literature indicate that severe losses of potential economic gain can possibly occur through use of a constrained index. It is concluded that constrained indexes should be avoided for economic genetic selection.     

Experimental mate replacement does not increase extra-pair paternity in tree swallows

Before the onset of female fertility, we removed 12 early-settling male tree swallows (Tachycineta bicolor) from their nest-boxes and mates, and allowed replacement males which had been floaters to settle with the original female residents. We predicted that females which had their original mate choice altered (experimentals) would be more likely to obtain extra-pair fertilizations than females which remained paired with their original, early-settling mates (controls). The proportion of females obtaining extra-pair fertilizations, however, did not differ between controls and experimentals, indicating that mating tactics of female tree swallows were unaffected by mate replacement. However, differences between early-settling and replacement males did exist. Replacement males had shorter wing chords than early-settling males, suggesting that they were younger. Moreover, a significantly greater proportion of replacement males than early-settling males were unringed and thus new to the study site. Our results suggest that if females are choosing males for good genes, then early-settling males are not superior in genetic quality to the smaller replacement males which had originally been floaters.     

Social organization in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

The evolution of female sexuality and mate selection in humans

Understanding female sexuality and mate choice is central to evolutionary scenarios of human social systems. Studies of female sexuality conducted by sex researchers in the United States since 1938 indicate that human females in general are concerned with their sexual well-being and are capable of sexual response parallel to that of males. Across cultures in general and in western societies in particular, females engage in extramarital affairs regularly, regardless of punishment by males or social disapproval. Families are usually concerned with marriage arrangements only insofar as those arrangements are economically or politically advantageous, but females most often have a voice in arranged marriages. Extended families also concentrate on a couple’s future reproduction rather than on sexual exclusivity. Although marriage for females is often compromised by male or family reproductive interests (which may not in fact differ from female interests), females appear to exercise their sexuality with more freedom than has previously been suggested. Notions of human females as pawns in the male reproductive game, or as traders of sex for male services, should be dispelled.     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

Experimental comparison of selection methods to improve a non-linear trait in Tribolium

Summary Five lines of Tribolium castaneum (flour beetle) were selected in each of three, replicates for biomass, using quadratic (Q) and linear (L) selection indices, direct selection for biomass (B) and indirect selection for each component trait, i.e. pupal weight (P) and family size (F), respectively, for five generations. The highest response in biomass was obtained in the Q line using an exact quadratic index in which the genetic value for each trait was based on information from both traits. The expected results were that Q selection would not exceed L selection while the L method would be superior with respect to direct selection for biomass (B line); P selection would be better and F selection worse than B selection. Selection was effective for pupal weight in the Q, L, B and P lines, but was not effective for family size, although the Q line increased slightly. The lines had no linear response for family size. Selection for biomass was influenced more by family size than by pupal weight. It is advantageous to include terms such as squares and products in the selection index, even though the response to selection may have been non-linear because of the low heritability of family size and an intermediate optimum for this component trait.