Fluctuating asymmetry in certain morphological traits in laboratory populations of Drosophila ananassae.

Fluctuating asymmetry (FA, subtle random deviations from perfect bilateral symmetry) is often used as a measure of developmental instability (DI), which results from perturbations in developmental pathways caused by genetic or environmental stressors. During the present study, we estimated FA in 5 morphological traits, viz. wing length (WL), wing to thorax ratio (W:T), sternopleural bristle number (SBN), sex-comb tooth number (SCTN), and ovariole number (ON) in 18 laboratory populations of Drosophila ananassae. FA levels of measured traits differed significantly among populations except for SBN (in males and females) and W:T ratio (in females). Positional fluctuating asymmetry (PFA), a sensitive measure of DI, also varied significantly among the populations for SBN in females and SCTN in males. Interestingly, both males and females were similar for nonsexual traits. However, when FA across all traits (sexual and nonsexual) was combined into a single composite index (CFA), significant differences were found for both populations and sexes. Males showed higher CFA values than females, suggesting that males are more prone to developmental perturbations. The magnitude of FA differed significantly among traits, being lowest for nonsexual traits (SBN, WL, W:T ratio) and highest for sexual traits (SCTN and ON). The trait size of sexual traits (SCTN and ON) was positively correlated with their asymmetry. The possible reasons for variation in FA both among traits and among populations, and the usefulness of FA as an indicator of developmental stress and phenotypic quality in D. ananassae are discussed.     

The relationship between sexual dimorphism in human faces and fluctuating asymmetry

Previous studies have found both support and lack of support for a positive relationship between masculinity and symmetry, two putative signs of mate quality, in male faces. We re-examined this relationship using an explicit measure of facial fluctuating asymmetry, as well as other measures of asymmetry, and measures of facial masculinity/femininity. We also used ratings of these traits for faces. Further, we examined the relationship between facial sexual dimorphism and body asymmetry. We found no significant correlations between facial masculinity and any of our measures of asymmetry or ratings of symmetry in males. Facial femininity was not consistently associated with facial symmetry in females, but was associated with body symmetry. Therefore, for females, but not males, facial femininity and body symmetry may reflect similar aspects of mate quality. We also examined the relationships between trait ratings and measurements. Our results provide validation of our ability to measure aspects of asymmetry that are perceived to be symmetrical, and aspects of sexual dimorphism that are perceived as feminine in females and masculine in males.     

Does urban poverty increase body fluctuating asymmetry?

Perturbations during development leave enduring signs on the adult body. Fluctuating asymmetry (FA) is a good bio-indicator of stress during ontogeny. The aim of this study is to determine the effect of urban poverty on the fluctuating asymmetry of young Turkish males. Young males from a lower socioeconomic group (N = 140, Mean age = 18.17 +/- 0.61) were selected from slum areas of Ankara, the capital of Turkey, where urban poverty is intense. An upper socioeconomic group, on the other hand, consisted of students from two private colleges and included children from some of the richest families in Turkey (N = 120, Mean age = 18.08 +/- 0.54). Eight anthropometric traits of all subjects were measured. Considering the seven measurements demonstrate ideal FA, the individuals living in poor areas of the city displayed higher FA. The discrepancy between the two groups was even greater for a measure of composite FA. In conclusion, poor living conditions in Ankara, where urban poverty is intense, adversely impact the developmental stability of young Turkish males.     

Do facial averageness and symmetry signal health?

We investigated whether the attractive facial traits of averageness and symmetry signal health, examining two aspects of signalling: whether these traits are perceived as healthy, and whether they provide accurate health information. In Study 1, we used morphing techniques to alter the averageness and symmetry of individual faces. Increases in both traits increased perceived health, and perceived health correlated negatively with rated distinctiveness (a converse measure of averageness) and positively with rated symmetry of the images. In Study 2, we examined whether these traits signal real, as well as perceived, health, in a sample of individuals for whom health scores, based on detailed medical records, were available. Perceived health correlated negatively with distinctiveness and asymmetry, replicating Study 1. Facial distinctiveness ratings of 17-year-olds were associated with poor childhood health in males, and poor current and adolescent health in females, although the last association was only marginally significant. Facial asymmetry of 17-year-olds was not associated with actual health. We discuss the implications of these results for a good genes account of facial preferences.     

Hand asymmetry does not relate to key life history traits in post-menopausal contemporary Finnish women

Associations between fluctuating asymmetry (FA, a putative marker of developmental instability, DI) and life history traits have received a great deal of attention in the non-human literature. However, the patterns found are very heterogeneous and generalizations are difficult to make. In humans, only a few studies have related FA to life histories and fitness. In this paper we study such relationships using hand FA and several key life history traits in 209 post-menopausal Finnish women born between 1946 and 1958. Asymmetry measurements were based on scans of the hands and the life histories of these women were collected using questionnaires. No significant associations were detected and trends were opposite to expectations. We did find evidence for directional asymmetry, as traits in the right hand were larger on average. This may be due to handedness, questioning the usefulness of hand FA as a measure of DI. We conclude that future studies in humans should carefully examine the usefulness of traits as measures of DI.     

Facial attractiveness signals different aspects of “quality” in women and men

We explored the relationships between facial attractiveness and several variables thought to be related to genotypic and phenotypic quality in humans (namely fluctuating asymmetry (FA), body mass index (BMI), health, age). To help resolve some controversy around previous studies, we used consistent measurement and statistical methods and relatively large samples of both female (n=94) and male (n=95) subjects (to be evaluated and measured), and female (n=226) and male (n=153) viewers (to rate attractiveness). We measured the asymmetry of 22 traits from three trait families (eight facial, nine body and five fingerprint traits) and constructed composite asymmetry indices of traits showing significant repeatability. Facial attractiveness was negatively related to an overall asymmetry index in both females and males, with almost identical slopes. Female facial attractiveness was best predicted by BMI and past health problems, whereas male facial attractiveness was best predicted by the socioeconomic status (SES) of their rearing environment. Composite FA indices accounted for a small (<4%) but usually significant percentage of the variation in facial attractiveness in both sexes, when factors related to asymmetry were controlled statistically. We conclude that, although facial attractiveness is negatively related to developmental instability (as measured by asymmetry), attractiveness also signals different aspects of “quality” in the two sexes, independent of FA.     

Human body odour, symmetry and attractiveness.

Several studies have found body and facial symmetry as well as attractiveness to be human mate choice criteria. These characteristics are presumed to signal developmental stability. Human body odour has been shown to influence female mate choice depending on the immune system, but the question of whether smell could signal general mate quality, as do other cues, was not addressed in previous studies. We compared ratings of body odour, attractiveness, and measurements of facial and body asymmetry of 16 male and 19 female subjects. Subjects wore a T-shirt for three consecutive nights under controlled conditions. Opposite-sex raters judged the odour of the T-shirts and another group evaluated portraits of the subjects for attractiveness. We measured seven bilateral traits of the subject's body to assess body asymmetry. Facial asymmetry was examined by distance measurements of portrait photographs. The results showed a significant positive correlation between facial attractiveness and sexiness of body odour for female subjects. We found positive relationships between body odour and attractiveness and negative ones between smell and body asymmetry for males only if female odour raters were in the most fertile phase of their menstrual cycle. The outcomes are discussed in the light of different male and female reproductive strategies.     

Sex and asymmetry in humans: what is the role of developmental instability?

Because asymmetric individuals are less attractive and may suffer from reduced fitness, bilateral asymmetry is widely believed to affect human sexual selection. Its evolutionary significance is based on the presumed relationship with developmental instability (DI). Yet, relationships between DI and bilateral asymmetry are often weak and possibly confounded by asymmetric mechanical loadings because of handedness. We related asymmetry in hands and faces to degrees of handedness and sexual behaviour in 100 humans. Handedness correlated to levels of asymmetry, thereby likely invalidating the use of asymmetry to estimate DI. For facial asymmetry, applying existing theoretical models refuted a link between asymmetry and DI. Explicit statistical modelling at the level of DI confirmed the absence of a link between DI and aspects of sexual behaviour. Nevertheless, asymmetries in both hands and face correlated significantly with sexual behaviour. We conclude that bilateral asymmetry per se, rather than its presumed link with DI, more likely relates to measures of human sexual behaviour. Because lateralization of behaviour appears widespread, evaluating the role of DI in evolution and ecology relies on a very critical selection of traits whose asymmetry can reliably reflect DI.     

Artificial selection reveals heritable variation for developmental instability

Fluctuating (nondirectional) asymmetry (FA) of bilaterally paired structures on a symmetrical organism is commonly used to assay the developmental instability (DI) caused by environmental or genetic factors. Although evidence for natural selection to reduce FA has been reported, evidence that FA (and by extension DI) is heritable is weak. We report the use of artificial selection to demonstrate heritable variation in the fluctuating asymmetry of interlandmark distances within the wing in an outbred population of Drosophila melanogaster. Our estimates for the heritability of FA range from 0% to 1% and result in estimates for the heritability of DI as large as 20%, comparable to values typical for life-history traits. These values indicate the existence of evolutionarily relevant genetic variation for DI and the effectiveness of selection for reduced FA suggests that natural selection has not fixed all the genetic variants that would improve developmental stability in these populations.     

Sexual selection, stabilizing selection and fluctuating asymmetry in two populations of pupfish (Cyprinodon pecosensis)

Fluctuating asymmetry of morphological traits is thought to reflect the capacity of a genotype to produce an integrated, functional phenotype. I tested three predictions. (1) In a polygynous breeding system, under intense sexual selection on males, breeding males should show greater symmetry in bilaterally symmetrical traits than non-breeding males or females. (2) If these traits are under stabilizing selection, highly symmetrical individuals also should be modal phenotypes, thus near the mean value for that trait, whereas individuals with increased asymmetry should represent marginal phenotypes, near the extremes of the distribution for that trait. (3) Differences in the intensity of sexual selection should be reflected in differences in the degree of fluctuating asymmetry between sexes among populations. I examined the relationship between male breeding status and the degree of fluctuating asymmetry of four bilaterally symmetrical- traits, preorbital and preopercular pores and pectoral and pelvic fin rays, in two populations of Pecos pupfish which differed in the intensity of sexual selection. These traits do not function in male-male competition or female choice, thus are not directly affected by sexual selection. In Mirror Lake breeding males, as a group, were most symmetrical for all four traits, while non-breeding males and females showed higher levels of fluctuating asymmetry. Similarly, symmetrical individuals also represented modal phenotypes for four traits (breeding males), and for three traits (non-breeding males and females). These patterns were not seen in the Lake Francis population, where breeding males were as asymmetrical as non-breeding males and females, and the degree of fluctuating symmetry did not differ between modal and marginal phenotypes for any of the four traits. When ecological conditions favour intense sexual selection, either through female choice, male-male competition, or both, breeding males represent the most fit phenotypes. Thus sexual selection reinforces the effects of stabilizing selection on characters that do not function as secondary sexual traits. However, when sexual selection is relaxed, differences between sexes disappear.     

Facial attractiveness,developmental stability,and fluctuating asymmetry

Despite robust cross-cultural reliability of human facial attractiveness ratings, research on facial attractiveness has only superficially addressed the connection between facial attractiveness and the history of sexual selection in Homo sapiens. There are reasons to believe that developmental stability and phenotypic quality are related. Recent studies of nonhuman animals indicate that developmental stability, measured as fluctuating asymmetry in generally bilateral symmetrical traits, is predictive of performance in sexual selection: Relatively symmetrical males are advantaged under sexual selection. This pattern is suggested by our study of facial attractiveness and fluctuating asymmetry in seven bilateral body traits in a student population. Overall, facial attractiveness negatively correlated with fluctuating asymmetry; the relation for men, but not for women, was statistically reliable. Possible confounding factors were controlled for in the analysis.     

Fluctuating asymmetry and age in children: evolutionary implications for the control of developmental stability

Fluctuating asymmetry (FA) is small deviations from perfect symmetry which reflect one component of fitness, i.e. developmental stability. There is accumulating evidence that low FA is important in inter- and intra-sexual selection in humans. However, there is little information on the pattern of FA in children. Data from cross-sectional studies of 680 participants from 2–18 years suggests that (1) both absolute and relative FA reduces with age until 10 years (2) there is an increase in FA in adolescents (11–15 years) with a peak at 13 years for males and 14 years for females (3) after 15 years there is reduction in FA which is maintained until 18 years. The importance of growth rate, metabolic maintenance and sex steroids on developmental stability is discussed.     

Fluctuating asymmetry of sexual and nonsexual traits in stalk‐eyed flies: a poor indicator of developmental stress and genetic quality

It has been proposed that females use fluctuating asymmetry (FA) in sexual ornaments to assess male quality. FA of sexual traits is predicted to show greater sensitivity to stress than FA of nonsexual traits, and to be heritable. We used a half‐sib mating design and manipulation of larval food environment to test these predictions on stalk‐eyed flies, Cyrtodiopsis dalmanni, in which females prefer males with larger eyespans. We measured size and FA of eyestalks and of two nonsexually selected characters, wing length and width. We found no evidence of an increase in FA under larval food stress in any of the individual traits, although trait size decreased under stress. We combined FA across traits into a single composite index, and found that males reared in the most benign larval environment had significantly higher composite FA than males reared on other media. There was no such effect in females. Heritability of FA was not significantly different from zero in any of the traits, in any of the environments, although trait sizes showed high heritability. We conclude that FA in sexual and nonsexual traits is a poor indicator of developmental stress and genetic quality.     

Are human preferences for facial symmetry focused on signals of developmental instability?

Humans find symmetrical faces more attractive than are asymmetricalfaces. Evolutionary psychologists claim that our preferencefor symmetry can be explained in the context of mate choicebecause symmetry is an honest indicator of the genetic qualityof potential mates. These arguments assume that asymmetry inhuman faces is fluctuating asymmetry (FA), because this formof asymmetry can be revealing of developmental instability.However, no study has yet examined the characteristics of facialasymmetry. Here we provide the first detailed study of the patternsof asymmetry in human faces. We measured asymmetry in 35 facialtraits. Although some traits had distributions characteristicof FA, many had distributions that characterize directionalasymmetry (DA); on average, both men and women had right hemi-facedominance. For DA traits we used deviations from the mean asymmetryas a measure of developmental instability. Our measures of asymmetryaccounted for a moderate proportion of the variance in perceivedsymmetry. Importantly, only FAs and random deviations from DAcontributed to people's perception of symmetry. DA was not importantin symmetry judgments. Faces rated as symmetrical were alsorated as attractive. Random deviations from DA were weakly relatedto women's attractiveness judgments of men's faces. DAs didnot influence attractiveness judgments. Our data suggest thatpeople focus on aspects of facial asymmetry that may be revealingof developmental instability. Further studies that isolate FAfrom other forms of asymmetry are required to accurately assessthe influence of developmental instability on the quality ofindividuals and its potential role in mate preferences.     

Fluctuating asymmetry in mice and rats: evaluation of the method

Fluctuating asymmetry, which reflects small, random deviations from symmetry in otherwise bilaterally symmetrical characters, may be used as an indicator of developmental instability in humans and farm animals, and it may also be applicable as a stress indicator. We intended to find a method to allow the use of fluctuating asymmetry as a stress indicator in laboratory animals. That method had to be reproducible and reliable. Furthermore, its applicability in laboratory animals would be improved if it was possible to obtain measurements on the skin surface that correlated with results obtained by measuring the skeleton directly. Seven traits in mice and five traits in rats were evaluated for their applicability for measuring fluctuating asymmetry in mice and rats. Two out of the seven traits, i.e. the width of the joint between the third metatarsal bone and the digital bone on the hind paw, and the length of the incisor tooth at the top, were found to be reliable and reproducible for detecting fluctuating asymmetry in mice as well as in rats. Three out of the seven traits, i.e. the width of the carpal bones, the width of the joint between the tibia and the tarsal bones, and the length of the incisor tooth at the bottom, did express fluctuating asymmetry, but showed a poor day-to-day reproducibility. If the day-to-day reproducibility could be increased, these three traits might also be suitable for measuring fluctuating asymmetry in mice and rats. The last two traits, i.e. the length of ulna and the length of calcaneus plus metatarsal bone i.v., measured both on the skin surface and directly on the bone, did not express fluctuating asymmetry, and had a poor day-to-day reproducibility. These two traits are not suitable for measuring fluctuating asymmetry in mice and rats.     

Larger swordtail females prefer asymmetrical males

Many organisms, including humans, find symmetry more attractive than asymmetry. Is this bias towards symmetry simply a by-product of their detection system? We examined female preference for symmetry of the pigment pattern vertical bars in the swordtail fishes Xiphophorus cortezi and Xiphophorus malinche. We found a relationship between preference for symmetry and female size, with larger and thus older females spending significantly more time with the asymmetrical video animation as compared to the symmetrical video animation. The preference for asymmetry we report demonstrates that even if females can detect symmetrical males better, this does not preclude subsequent selection on females to prefer symmetrical or asymmetrical males. In addition, because the preference was correlated with female size, past studies may have missed preference for either asymmetry and/or symmetry by not examining the relationship between female preference and size/age or by measuring a limited size/age distribution of females. In both of the species of swordtail fishes examined, a high proportion of males are asymmetrical by more than one bar. We suggest that female preference may be maintaining fluctuating asymmetries in these fishes.     

Female health,attractiveness, and desirability for relationships: Role of breast asymmetry and waist-to-hip ratio

Fluctuating asymmetry (FA) of ordinarily bilaterally symmetrical traits in humans has been proposed to indicate developmental anomaly. Recent research has shown that individuals with minimal FA are judged to be attractive, and preferred as sexual partners (Thornhill and Gangestad 1993). Waist-to-hip ratios (WHR) have been also shown to reflect health and reproductive capability of woman and those with low WHRs are judged more attractive and healthy (Singh 1993a,b). The present study examines the relative roles of WHR and FA in female breasts in judgments of female attractiveness, health, and desirability for short-term and long-term relationships. Male college students were asked to judge attractiveness of female figures that differed in WHR (high and low) and breast symmetry (none, low, or high). In the first test, paired comparison method was used in which each figure was paired one at a time with all other figures. In the second test, subjects examined all figures simultaneously, estimated their age, and rated each figure for attractiveness, feminine looks, health, overall degree of body symmetry, and willingness to engage in short- and long-term relationship. Results from both tests show that figures with low WHRs were judged to be more attractive than figures with high WHRs, regardless of their degree of breast asymmetry. The figure with low WHR and symmetrical breasts was judged to be most attractive and youngest of all other figures. It appears that men use both WHR and breast asymmetry in judging attractiveness and being willing to develop romantic relationships. It is proposed that WHR and breast asymmetry may signal different aspects of overall female mate quality.     

Male facial attractiveness and masculinity may provide sex‐ and culture‐independent cues to semen quality

Phenotype‐linked fertility hypothesis (PLFH) predicts that male secondary sexual traits reveal honest information about male fertilization ability. However, PLFH has rarely been studied in humans. The aim of the present study was to test PLFH in humans and to investigate whether potential ability to select fertile partners is independent of sex or cultural background. We found that on the contrary to the hypothesis, facial masculinity was negatively associated with semen quality. As increased levels of testosterone have been demonstrated to impair sperm production, this finding may indicate a trade‐off between investments in secondary sexual signalling (i.e. facial masculinity) and fertility or status‐dependent differences in investments in semen quality. In both sexes and nationalities (Spanish and Colombian), ranked male facial attractiveness predicted male semen quality. However, Spanish males and females estimated facial images generally more attractive (gave higher ranks) than Colombian raters, and in both nationalities, males gave higher ranks than females. This suggests that male facial cues may provide culture‐ and sex‐independent information about male fertility. However, our results also indicate that humans may be more sensitive to facial attractiveness cues within their own populations and also that males may generally overestimate the attractiveness of other men to females.     

先天性心脏病患者皮纹波动不对称性的研究

本文采用随机整体抽样的方法分析了先天性心脏病患者129例(男性59例, 女性70例) 和正常对照人群133例(男性69例, 女性64例) 13项皮纹波动不对称性(Fluctuating asymmetry, FA)的分布特征。结果表明: (1)先天性心脏病患者组与正常对照组在13项皮纹波动不对称性指标中均未出现显著性别差异; (2)先天性心脏病患者组与正常对照组在FAⅥ(P<0.05)和FAⅦ(P<0.01)两项有显著性差异, 表现为患者组明显增高, 提示先天性心脏病患者在胚胎发育早期易受到环境因素影响, 具有较高的发育不稳定性。     

Symmetry: attractive not only to females

Fluctuating asymmetry in bilateral traits has been proposed to reflect aspects of individual quality, and has hence been suggested to act as a cue in mate choice. Since sexual selection generally acts more strongly on males, numerous studies have focused on female preferences for symmetrical partners, while very little is known about potential male symmetry preferences. In the present experiment, we tested whether bluethroat males are sensitive to symmetry in an artificial ornament. Using different combinations of blue and orange leg bands, females were made symmetrical or asymmetrical. In outdoor aviaries placed in breeding habitat, males were allowed to choose between a symmetrical and an asymmetrical female. We found that males associated more with symmetrical than asymmetrical females, indicating a preference for symmetry. The magnitude of the symmetry preference of bluethroat males was similar to that of females of the same species.