A polymorphism in female preference for a polymorphic male trait in the swordtail fish Xiphophorus cortezi

Xiphophorus cortezi males are polymorphic for the pigment pattern vertical bars. In this study, we determined whether X. cortezi females are polymorphic in their preference for this trait by examining both within- and between-individual variation in female preference. There was significantly more variation in female preference within than between individuals using both video animations and live males as stimuli; repeatability measures were 0.86 and 0.5, respectively. Some females had a strong preference for males with bars, some for males without bars, and some had either a weak preference or no preference at all. We also found a significant difference in the strength of preference for bars between females with and without bars, suggesting the potential for a genetic correlation between preference and trait. Finally, we examined female preference for bar symmetry in the same females tested for preference for bars. We detected a significant preference for bar symmetry in the population of females as a whole, and a positive relationship between the strength of this preference and a preference for the presence/absence of bars among the subset of females that preferred ‘no bars’. We discuss these results in light the possibility that these two preferences are interrelated. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Large body size in the pygmy swordtail Xiphophorus pygmaeus

Fishes in the genus Xiphophorus (swordtails and platyfishes) are well known for the influence of the pituitary ( P ) locus on variation of male size at maturity both within and among species. We report the discovery of large male size (>29 mm SL) in several populations of the swordtail X. pygmaeus , a species previously thought to consist of only small males (<29 mm SL). Large size is geographically restricted, and average male size varies significantly by site and year sampled in a pattern suggesting a recent origin and slow spreading of the large male phenotype. However, large male size is not strongly paternally inherited in this species, as it is in its two closest relatives, X. nigrensis and X. multilineatus , showing that large size does not result from the same genetic ( P locus) mechanism. Large X. pygmaeus males do not court, can exhibit the gold morph, do not possess swords and have slender body shape. In these traits they resemble small conspecific males and small males of X. nigrensis and X. multilineatus rather than large males of these latter two species. This shows that correlations between morphological and behavioural traits that occur in X. nigrensis and X. multilineatus are absent in X. pygmaeus.     

Defining Vertical Bars in Relation to Female Preference in the Swordtail Fish Xiphophorus cortezi (Cyprinodontiformes, Poeciliidae)

Defining a male trait in relation to female preference is an important step towards determining the role that female preference has played in the evolution of a male trait. The pigment pattern 'vertical bars' is a complex male trait that functions as a sexual signal in swordtail fishes. A previous study suggested that X. cortezi females had a preference for males with more vertical bars. In that study, however, bar number had been reduced on the stimulus males by removing the most posterior bars, which changed the distance over which the bars spanned, in addition to changing bar number. To determine whether X. cortezi females have a preference for greater bar span and/or number of bars, we tested for female preference in two experiments. First, we gave females a choice between males with a greater bar span or a smaller bar span, holding bar number and total pigmented area constant. Females spent significantly more time with the males with the smaller bar span. This result, together with results from the previous study, suggest that females do not assess bar span alone, but possibly a composite component of the bars, such as bar frequency (number of bars/bar span). Secondly, we gave females a choice between six thinner bars or one wider bar of equal pigmented area. The six-bar treatment had a greater bar span in addition to more bars. We detected no preference for either treatment. These results suggest that the previously detected preference for more bars may actually reflect a preference for greater pigmented area. Finally, we present data on natural variation in the bars for X. cortezi and use principal components analysis to provide a composite definition of this trait.     

Vertical bars on male Xiphophorus multilineatus: a signal that deters rival males and attracts females

We examined the function of the vertical bar pattern on maleswordtails (Xiphophorus multilinneatus) as a signal in bothmale-male competition and female choice. This pattern had previouslybeen described as an aggressive signal because males intensifiedthe bars during male-male encounters in the laboratory. Ourfield observations supported this observation and also showedthat bars intensified when males courted females. The intensityof bars was correlated with access to females in the field.Within the size range of males that have bars, however, neitherbar number nor male size appeared to influence access to females.We used freeze-branding to remove the bars from males in thelaboratory so that we could control for characters correlatedwith bar intensity, and tested males and females separatelyso that we could separate the influence of these two componentsof sexual selection. We compared the responses of males andfemales to males that had their bars removed and control malesfreeze-branded between the bars. Test males responded more aggressivelyto males without bars as compared to control males. In addition,females showed a preference for control males over males thathad their bars removed. These results suggest that the barsmay function as a signal that deters rival males and attractsfemales.     

A Technique for the Temporary Application and Augmentation of Pigment Patterns in Fish

Secondary sexual characteristics, such as pigment patterns in male fish, have significant roles in female mating preferences. Vertical body-bar pigmentation, one such characteristic, is found in a variety of Xiphophorus species. Prior investigations have revealed that these bars function as a signal attracting females and deterring rival males in at least two species. Furthermore, it has been shown that X. cortezi females prefer vertical bar symmetry as well as more bars in conspecific males. Because of the shortcomings of prior techniques used for pigment elimination and permanent color marking, a temporary method for augmenting pigment patterns with an artificial dye was developed. This technique was tested using the known preference that female X. cortezi have for more bars in conspecific males. The results corroborate previous findings as female X. cortezi preferred males with more artificial vertical bars in comparison with males with fewer. Therefore, the method of applying temporary, artificial pigment patterns using an antiseptic dye would appear to be an effective way to control and manipulate secondary sexual pigmentation for behavioral investigations in fish.     

Female preference for trait symmetry in addition to trait size in swordtail fish

Interest in fluctuating asymmetries, random deviations from perfect bilateral symmetry, has spread to studies of sexual selection because of the intriguing idea that females could use the degree of asymmetry of a male trait to assess the genetic quality of potential mates. The evidence that females prefer males with symmetrical sexual signals, however, remains controversial. A problem that applies to most previous studies is that preference for trait size can be misinterpreted as preference for symmetry, even when overall trait size is held constant, if females assess trait size by the largest minimum on one side. If overall trait size is equal between males, the asymmetrical males will have the maximum and minimum trait size, and so preference to mate with symmetrical males could actually reflect a preference to avoid males with the minimum trait on one side. Xiphophorus cortezi females preferred males with symmetrical bar numbers when the minimum number of bars was held constant. The strength of female preference for the symmetrical males was negatively correlated with the strength of preference the same females had for bar number. These results clearly demonstrate that females preferred trait symmetry in addition to trait size.     

Sex-specific effects of gonadal steroids on conspecific odor preference in the rat

Effects of gonadal steroids on conspecific odor preference for either (1) sexually active male or active female, (2) sexually active or gonadectomized (gdx) males, (3) sexually active or gdx females, and (4) gdx males or gdx females were determined in male and female rats in a three-chamber apparatus. For the first test, gdx females were made sexually active by treatments with estradiol benzoate (EB) and progesterone (P), and sexually active males were selected by prior screening. Sexually active males and females preferred opposite-sex odor over same-sex odor. Odor of sexually active opposite-sex conspecifics was preferred over that of inactive ones. Immediately after the completion of the first test, sexually active males were gdx and females were left without hormonal treatment. Second and third tests were carried out 2 and 5 weeks after the first test. In the second test, gdx males preferred odor of sexually active males rather than that of receptive females (male-directed preference); in the third test, both males and females showed no preference when tested with four stimulus pairs. The final tests were carried out in gdx males with EB and P, and gdx females with 2-week exposure to testosterone (T). Males with EB and P showed a male-directed preference again, whereas T-treated females kept their own female preference. Injection of EB alone to gdx males did not induce any preference. The present study clearly demonstrated sex difference in conspecific odor preference. Although both male and female preferences depend on their circulating sex steroids, the direction of male preference is more susceptible to their hormonal states, compared to that of females.     

Female preference for dominant males in the Mexican mojarra cichlid fish,Cichlasoma istlanum

It is widely assumed that female preference and male competition operate simultaneously during sexual selection. Dominance is likely an honest indicator of male quality, and females can identify and choose the dominant male to reproduce with individuals with greater competitive abilities, thus improving the quality and competitiveness of their offspring. In this context, few studies have investigated female preference in relation to male fighting ability. The Mexican mojarra, Cichlasoma istlanum, is a cichlid species native to the Balsas River basin. It is territorial during reproduction and provides parental care. Males commonly engage in territorial defence, whereas females care directly for offspring. This study examined whether females prefer dominant males that exhibit more aggressive behaviour. The authors conducted experiments using groups of two males and one female to test competitive ability in males and female preference. They also quantified the time during which the female associated with the dominant male and the subordinate male after observing the outcome of a fight between the two males. They found that Mexican mojarra females preferred dominant males and that the time females spent associating with males was positively related with their aggressive behaviours during competition. These results indicate that dominant males were more attractive than subordinate males to female Mexican mojarra. The relationship between female preference and male dominance in the Mexican mojarra demonstrates the importance of male competitive ability for future parental care in reproduction.     

Variation in female guppy preference for male olfactory and visual traits

Animals often use different sensory systems to assess different sexually selected signals from potential mates. However, the relative importance of different signals on mate choice is not well understood in many animal species. In this study, we examined the relative importance of male olfactory and visual cues on female preference in the guppy Poecilia reticulata. We used digitally modified male images to standardize visual stimuli. We found that, regardless of whether females were presented without male visual stimuli or with identical male visual stimuli, they preferred stimuli with the odor of males to those without. However, when females were allowed to choose between dull male visual stimuli with male odor, and brightly colored male visual stimuli without male odor, there was no clear preference for either. Some females preferred the dull male visual stimuli with male odor, whereas some other females preferred the brightly colored male visual stimuli without male odor. These results indicate that the relative importance of olfactory and visual cues in female mate preference varied between individuals.     

Genetics, experience, and host-plant preference in Eurosta solidaginis: implications for host shifts and speciation

Host-associated mating is crucial in maintaining the partial reproductive isolation between the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and S. gigantea. (We refer to flies reared from S. gigantea as gigantea flies and those reared from S. altissima as altissima flies.) We measured the host preference of males and females of both host races, F1 hybrids between the host races, F2, and backcrosses to both host races. Male and female altissima flies and female gigantea flies had high host fidelity, whereas male gigantea flies had low host fidelity. This result suggests that there may be gene flow between the host races due to nonassortative mating that occurs when male gigantea mate with altissima females on S. altissima. This indicates assortative-mating mechanisms in addition to host-associated mating are required to produce the partial reproductive isolation between the host races that has been observed. Nongenetic factors had no influence on host preference. Larval conditioning did not influence host preference: reciprocal F1 hybrids reared in S. altissima and S. gigantea both preferred S. gigantea. Adult experience had no impact on host preference: females preferred their natal host plant regardless of which host they encountered first as an adult. The hypothesis that maternal effects influence preferences was rejected because male and female flies did not show a consistent preference for the host plant of their mother. We also found no evidence that preference was a sex-linked trait because F1 and backcrosses to the host races with different combinations of X chromosomes from the two host races preferred S. gigantea. Our results indicate that host preference is not determined by a large number of genes because preference of hybrids did not correspond to the proportion of the genome derived from each host race. The strength of the ovipuncture preference for S. gigantea by gigantea females, the females of both reciprocal F1 hybrids, the backcross to gigantea, and F2s indicates that preference is inherited nonadditively at a limited number of loci. The F1 female hybrids, however, had a weaker host preference for S. gigantea than the pure gigantea host race, indicating that there may be incomplete dominance or modifier loci. Males had different host preference patterns than females, with individual male gigantea and male F1 hybrids usually exhibiting preference exclusively for S. gigantea or S. altissima. One hypothesis explaining the difference in host preference between males and females is that the same gene influences both female and male host preference, but it is a sex-influenced gene. Thus, males carrying the gene for S. gigantea preference have an intermediate host preference, whereas females have a strong host preference to S. gigantea. In summary, we found that the host preference that produces host-associated mating is inherited nonadditively at a relatively small number of loci on autosomal genes. This mode of inheritance meets the assumptions of models of sympatric speciation, indicating that the host races could have evolved in sympatry.     

Female preference for preferred males is reversed under low oxygen conditions in the common goby (Pomatoschistus microps)

Female preference for males that already have eggs in theirnest has been reported in many fish species. The presence ofeggs may provide a cue for copying the choice of previous femalesand may indicate that high-quality care will be available. Consistentwith a review of 13 studies, we found that female common gobies(Pomatoschistus microps) preferred to spawn with males thathad already been chosen by females and whose nests therefore alreadycontained eggs. However, this preference was reversed underconditions of low dissolved oxygen. We would not expect thisreversal if the second female were using eggs as a signal ofmale genetic attractiveness to other females unless the benefitswere outweighed by direct selection. The reversal also couldnot be explained by differences in active courtship by males,as courtship rates did not differ under low oxygen between maleswith or without eggs. Low oxygen conditions corresponded witha nearly threefold increase in male ventilation of eggs anda reduction in time spent near a selecting female. The reversalis therefore most likely due to females avoiding males thatwould be unable to meet the demands of care of a second clutchunder low oxygen conditions. Thus, an abiotic feature of theenvironment reveals plasticity of female choice, consistentwith hypothesized changes in benefits of mating with preferredmales.     

Size-dependent mating preference of the male fiddler crab Austruca perplexa

In many animals, females prefer large males to small males, which allow large males to be choosier than small males when selecting a mate. We investigated the courtship intensity of small- and large-sized male fiddler crabs (Austruca perplexa) by examining their claw-waving rates (waves/min) towards small- and large-sized females. We found that large males showed a greater preference for large females by producing more waves/min towards them, whereas small males did not show any apparent preference for either large or small females. Moreover, the waving rate of large males was positively correlated with female size, but there was no correlation between waving rate and female size in small males. These results indicate that large males in a population become choosier and show strong mate choice, which is most likely due to their greater preference among females.     

Divergent female mating preference in a clonal fish

In asexual all-female species, such as the Amazon molly, Poecilia formosa, that depend on sperm from “host males” only to trigger embryogenesis, mate choice does not provide any indirect, genetic benefits to the choosing female, although direct benefits are possible. Asexual species are thought to have a low evolutionary potential or evolvability due to the absence of meiotic recombination. Hence, theory predicts that mating preferences in P. formosa for male ornaments—if existent—should resemble those of females of the two parental species (Poecilia latipinna and Poecilia mexicana) involved in the natural hybridization that gave rise to the asexual P. formosa. When examining the female preference for dummy males with or without black vertical bars in the two parental species and in two lineages of P. formosa, a preference was detected in P. latipinna, but not in P. mexicana females. Interestingly, P. formosa living syntopic with P. latipinna also preferred striped males, while others living syntopic with P. mexicana preferred non-striped males. The evolutionary significance of this phenomenon remains largely unexplained, but it might indicate the evolution of mating preferences in a species with low evolutionary potential. Possible mechanisms include introgression and mitotic gene conversion. Females might use male coloration as indicator mechanisms for male traits that matter in terms of direct benefits.     

Experience Plays a Role in Female Preference for Symmetry in the Swordtail Fish Xiphophorus malinche

Variation in female mating preferences was previously detected in wild-caught Xiphophorus cortezi and Xiphophorus malinche females: smaller (presumed younger) females preferred symmetrical males, while larger (presumed older) females preferred asymmetrical males. We examined the influence of experience on this variation in female preference by determining if X. malinche females would express a preference for symmetry as virgins, shift their preferences for bar symmetry as they got larger (older) and if experience with males of different bar number symmetry could explain the variation in female preference previously detected. Virgin females exhibited no preference for vertical bar number symmetry when tested in the young- or old-age classes. However, young virgins spent more time with the opposite treatment in the second when compared with first test, indicating an ability to detect the difference between symmetry and asymmetry, and potentially a preference to mate with multiple males. When females were reared in one of three treatments, housed with symmetrical, barless or both symmetrical and asymmetrical males, we detected both a treatment and tank effect on strength of preference for symmetry, suggesting that barring pattern and some other aspect of the social environment influenced the development of this mating preference. Finally, we detected no effect of age class on mean strength of preference for symmetry; however, there was a statistically different relationship between female size and strength of preference for symmetry across the two age classes, suggesting that the preference function for symmetry may not be linear in relation to female size.     

Male competition reverses female preference for male chemical cues

Females must choose among potential mates with different phenotypes in a variety of social contexts. Many male traits are inherent and unchanging, but others are labile to social context. Competition, for example, can cause physiological changes that reflect recent wins and losses that fluctuate throughout time. We may expect females to respond differently to males depending on the outcome of their most recent fight. In Bolitotherus cornutus (forked fungus beetles), males compete for access to females, but copulation requires female cooperation. In this study, we use behavioral trials to determine whether females use chemical cues to differentiate between males and whether the outcome of recent male competition alters female preference. We measured female association time with chemical cues of two size‐matched males both before and after male–male competition. Females in our study preferred to associate with future losers before males interacted, but changed their preference for realized winners following male competitive interactions. Our study provides the first evidence of change in female preference based solely on the outcome of male–male competition.     

Evolution of female preference for younger males

Previous theoretical work has suggested that females should prefer to mate with older males, as older males should have higher fitness than the average fitness of the cohort into which they were born. However, studies in humans and model organisms have shown that as males age, they accumulate deleterious mutations in their germ-line at an ever-increasing rate, thereby reducing the quality of genes passed on to the next generation. Thus, older males may produce relatively poor-quality offspring. To better understand how male age influences female mate preference and offspring quality, we used a genetic algorithm model to study the effect of age-related increases in male genetic load on female mate preference. When we incorporate age-related increases in mutation load in males into our model, we find that females evolve a preference for younger males. Females in this model could determine a male's age, but not his inherited genotype nor his mutation load. Nevertheless, females evolved age-preferences that led them to mate with males that had low mutation loads, but showed no preference for males with respect to their somatic quality. These results suggest that germ-line quality, rather than somatic quality, should be the focus of female preference in good genes models.     

Context‐dependent female preference for multiple ornaments in the bearded reedling

While it is well established that females prefer to mate with well‐ornamented males, the influence of perceptive and cognitive processes on the expression of female mate choice is still poorly known. It has been suggested that the female perception of a male's attractiveness is not absolute, but depends on the other males with which he is compared that have been previously encountered (comparative evaluation). We investigated whether mate preference in bearded reedlings (Panurus biarmicus) is dependent on or independent of social context in relation to two different traits: beard and tail lengths. Each female had a choice between two to three males with different modifications of beard and tail. For each female, three different experiments were conducted (one binary and two trinary tests). We found that when females are presented with options that vary antagonistically with respect to two ornaments (binary test), some individuals prefer one trait while others the other trait. This indicates that in our bearded reedlings population exists a mate preference polymorphisms. Moreover, we found that the presence of a third stimulus, irrespective of the initial preference, reduced the strength of the initial preference – what we can call a “preference dilution effect.” Our results suggest that the female's choice may be constrained by her cognitive abilities when she is simultaneously presented with several options varying for two uncorrelated traits.     

Genotype-by-environment interactions for female preference

Sexual selection is responsible for many of the most spectacular displays in nature, and female preference for certain males is central to much of this. However, female preference is relatively poorly understood, particularly the relative importance of a female's genes, the environment and their interaction on her preference. We investigated preference in a no-choice design using Drosophila melanogaster iso-female lines and find that there are genotype-by-environment interactions for female preference. Whereas the choosiness of some female genotypes differed little across environments, that of others differed greatly, so that the choosiness rank of females in one environment did not necessarily predict their rank in another. Furthermore, the genetic variance underlying preference also varied across environments. These findings have important consequences for the evolution of female preference and the male sexual traits preference targets.     

Adult partner preference and sexual behavior of male rats affected by perinatal endocrine manipulations

Intact adult male rats, in which aromatization of testosterone to estradiol was prevented pre- and/or neonatally by ATD (1,4,6-androstatriene-3,17-dione), were repeatedly tested for partner preference behavior (choice: estrous female vs active male). In consecutive tests increasing preference scores for the female were found. Neonatal ATD males showed significantly lower preference scores for an estrous female than controls or prenatal ATD males. Prenatal ATD caused preference scores only slightly lower than those of controls. Ejaculation frequencies were markedly reduced or even absent in neonatal ATD males. Prenatal ATD treatment only had no or a moderately lowering effect on ejaculation frequency. Lordosis behavior of adult intact males was more facilitated following neonatal ATD treatment than following prenatal ATD treatment. In a number of tests the serotonergic drug 8-OH-DPAT was injected prior to testing for sexual partner preference and copulatory behavior. DPAT significantly increased preference for an estrous female in all groups of males when interaction was possible, but had no effect when sexual interaction was prevented by wire mesh. DPAT was able to increase the number of ejaculators in nonejaculating groups (i.e., perinatally ATD-treated males). "Premature ejaculations," i.e., ejaculations with the first intromission, were frequently observed with DPAT treatment in all groups of males. In conclusion, the availability of neonatal estrogen (derived from testosterone) organizes, at least partially, the preference for an estrous female normally shown by adult male rats. The lack of neonatal estrogen causes males to be less masculinized, both in partner preference behavior and ejaculatory behavior, and less defeminized in lordosis behavior.(ABSTRACT TRUNCATED AT 250 WORDS)     

Genetic analysis of female preference functions as function-valued traits

The genetic analysis of female preferences has been seen as a particularly challenging empirical endeavor because of difficulties in generating suitable preference metrics in experiments large enough to adequately characterize variation. In this article, we take an alternative approach, treating female preference as a function-valued trait and exploiting random-coefficient models to characterize the genetic basis of female preference without measuring preference functions in each individual. Applying this approach to Drosophila bunnanda, in which females assess males through a multivariate contact pheromone system, we gain three valuable insights into the genetic basis of female preference functions. First, most genetic variation was attributable to one eigenfunction, suggesting shared genetic control of preferences for nine male pheromones. Second, genetic variance in female preference functions was not associated with genetic variance in the pheromones, implying that genetic variation in female preference did not maintain genetic variation in male traits. Finally, breeding values for female preference functions were skewed away from the direction of selection on the male traits, suggesting directional selection on female preferences. The genetic analysis of female preference functions as function-valued traits offers a robust statistical framework for investigations of female preference, in addition to alleviating some experimental difficulties associated with estimating variation in preference functions.