Effects of maternal body morphology, morning sickness, gestational diabetes and hypertension on fluctuating asymmetry in young women

The magnitude of fluctuating asymmetry (FA)—an indicator of genetic and phenotypic quality—can be affected by genetic perturbations, environmental stressors, and maternal effect (maternal age, diseases, dietary deficiency). Maternal effect on human FA has been typically investigated in newborns or very young children. There are no studies investigating whether maternal effect can disrupt developmental mechanisms responsible for the secondary sexual traits that are manifested at adulthood under the influence of steroid hormones. We investigated the effect of maternal degree of obesity, gestational diabetes and hypertension, and morning sickness on the magnitude of FA in nonsexual traits as well as asymmetric thigh circumference—a sexually differentiated trait—in adult daughters. Results revealed that gestational diabetes and hypertension and maternal obesity are positively associated with FA in nonsexual traits. FA in nonsexual traits was not associated with morning sickness; however, the FA in the sexual trait (thigh circumference) was positively related to third-trimester morning sickness. Fluctuating asymmetries of nonsexual traits and thigh circumference were significantly correlated. This preliminary study demonstrates a maternal effect on adult daughters' developmental instability as measured by sexual and nonsexual traits.     

Variation of anthropometric traits in normal newborns]

Variation of the set of anthropometric traits was studied in 302 normal singleton newborns (174 boys, 128 girls). Birth order has a significant effect on birth weight, body length and circumference of head and breast. The number of minor deviations from development per newborn (stigma) is negatively correlated with body weight and size. The number of stigma is significantly higher in a group of newborns deviated in normal body proportions. The index of fluctuating asymmetry is negatively correlated with body weight and size.     

Ear asymmetry and left-side cradling

Women and girls tend to cradle infants and dolls on the left side of the body. Left-sided cradling is found in chimpanzees and gorillas, is cross-cultural and present in historical works of art, and is transmitted down the human maternal line. One explanation for the left-cradling tendency is that it facilitates the flow of affective information from the infant via the left ear and eye to the center for emotional decoding, that is, the right hemisphere of the mother. We show that the developmental stability of the ear, as measured by ear asymmetry, is negatively correlated with the left-sided cradling tendency. Left-cradling English women holding infants and Jamaican girls holding dolls had a strong tendency to show lower ear asymmetry than right cradlers, whereas no such relationship was found in boys nor for various measures of asymmetry of the hand, with the possible exception of the wrist in Jamaican girls. Degree of handedness, as measured by the Annett peg-moving test, did not predict cradling preference in the Jamaican children, and the relationship between ear asymmetry and cradling preference was independent of hand preference. Our results suggest that developmental instability of the ear (including the pinna, external auditory meatus, and middle ear) may interfere with the flow of affective information to the right hemisphere. Ear asymmetry also showed evidence of strong maternal but not paternal transmission. It is suggested that between-individual variation of in utero stress may explain patterns of maternal transmission of lateral cradling tendencies.     

Development of fluctuating asymmetry in tail feathers of the barn swallow Hirundo rustica

Fluctuating asymmetry represents usually small, random deviations from symmetry in bilateral morphological characters. The ontogeny of asymmetry in morphological characters may reveal information about developmental processes in a general sense. I studied the development of fluctuating asymmetry in feather characters of the barn swallow Hirundo rustica, that are developed repeatedly during the single annual moult, with the following results. First, the side developing a larger feather was found to be partially biased, as demonstrated by one side consistently developing a larger feather under natural and experimentally induced growth episode events. Second, asymmetric feathers were found to consist of asymmetric daily growth increments, and the size of the increments developing under different environmental conditions were positively correlated. Third, fluctuating asymmetries of feathers developing under different environmental conditions were positively correlated, although the level of asymmetry was larger under adverse environmental conditions. Fourth, individual asymmetries in tail length and growth bar length were unrelated to the duration of the developmental period, although late growth increments were smaller and more symmetric than early increments. These observations suggest that fluctuating asymmetry partially arises as a consequence of a random bias in the feather follicles and differences in environmental conditions during ontogeny of feathers.     

Genetic and phenotypic correlation between fluctuating asymmetry and two measurements of fear and stress in chickens

The purpose of the present study was to estimate the genetic and phenotypic correlation between fluctuating asymmetry and two measurements of fear and stress in chickens which had not deliberately stressed in any way, using the restricted maximum likelihood procedure. A total of 1073 36-week-old birds from two generations with complete pedigree of the Quail Castellana breed was used. Fluctuating asymmetry of several traits (leg, wing, and feather lengths, and ear-lobe and wattle areas), tonic immobility duration (indicator of fear), and heterophil to lymphocyte ratio (indicator of stress) were measured. The estimated genetic relationship between relative fluctuating asymmetry for the different traits and tonic immobility tended to be positive, that between the combined relative asymmetry of all traits and tonic immobility being near to +1; no significant phenotypic relationship was found between relative fluctuating asymmetry and tonic immobility. The genetic relationship between relative fluctuating asymmetry and heterophil to lymphocyte ratio was not consistent across the traits, ranging from +1 to −1, although the genetic correlation between the combined relative asymmetry and the heterophil to lymphocyte ratio was near to +1 too; no significant phenotypic relationship was found between relative fluctuating asymmetry and heterophil to lymphocyte ratio either. Relative fluctuating asymmetry and body weight were genetically negatively correlated for leg length and ear-lobe area but positively for feather length; the genetic correlation between the combined relative asymmetry and the body weight being near to −1; phenotypic relationships were not significantly different from zero. A significant negative genetic correlation between tonic immobility and heterophil to lymphocyte ratio was found, although the phenotypic association between these two measurements was zero. Phenotypic correlations always near to zero suggest that fluctuating asymmetry was not associated with duration of tonic immobility and heterophil to lymphocyte ratio in birds that have not been deliberately stressed.     

Fluctuating asymmetry and sexual selection

Fluctuating asymmetry occurs when an individual is unable to undergo identical development on both sides of a bilaterally symmetrical trait. Fluctuating asymmetry measures the sensitivity of development to a wide array of genetic and environmental stresses. We propose that fluctuating asymmetry is used in many signalling contexts for assessment of an individual's ability to cope with its environment. We hypothesize that fluctuating asymmetry is used in sexual selection, both in fighting and mate choice, and in competition for access to resources. Evidence is reviewed showing that the patterns of fluctuating asymmetry in secondary sexual characters differ from those seen in other morphological traits. Secondary sexual characters show much higher levels of fluctuating asymmetry. Also, there is often a negative relationship between fluctuating asymmetry and the absolute size of ornaments, whereas the relationship is typically U-shaped in other morphological traits. The common negative relationship between fluctuating asymmetry and ornament size suggests that many ornaments reliably reflect individual quality.     

Parent Barn Swallow fluctuating asymmetry and offspring quality

Fluctuating asymmetry may impair locomotion but may also reflect intrinsic phenotypic quality. I tested whether fluctuating asymmetry of adult Barn Swallows Hirundo rustica negatively influenced offspring quality, by estimating the relationship between parental asymmetry and offspring size, condition and immunocompetence during three breeding seasons. Controlling for timing of breeding, brood size and the size of a secondary sexual character (tail length), wing and outermost tail feather asymmetry of male and female parents was not significantly correlated with offspring size, condition and immunocompetence. This was the case in spite of clear differences in nestling quality among years. In addition, parents with extreme asymmetries due to tail feather damage (not representing fluctuating asymmetry) did not have nestlings of lower quality than parents with undamaged tail feathers. These results indicate that there is only a weak relation between parental asymmetry and offspring quality.     

The Symmetry of Children’s Knees Is Linked to Their Adult Sprinting Speed and Their Willingness to Sprint in a Long-Term Jamaican Study

Jamaican athletes are prominent in sprint running but the reasons for their success are not clear. Here we consider the possibility that symmetry, particularly symmetry of the legs, in Jamaican children is linked to high sprinting speed in adults. Our study population was a cohort of 288 rural children, mean age 8.2 (±1 SD = 1.7) years in 1996. Symmetry was measured in 1996 and 2006 from the fluctuating asymmetry (FA) of three lower-body traits and we constructed a lower body composite FA trait (Comp lb-FA). In 2010 we measured sprinting speed (for 90 m and 180 m races) in participants recruited from our original cohort. There were 163 untrained adults in our sample. We found: (i) high Comp lb and knee symmetry in 1996 and 2006 were linked to fast sprinting times in our 2010 runners and (ii) our sample of sprinters appears to have self-selected for greater symmetry. We conclude that high knee symmetry in childhood is linked to an ability to sprint fast in adult Jamaicans as well as a readiness to sprint.     

Fluctuating asymmetry and aggression in boys

Fluctuating asymmetry (FA) is small deviations from perfect symmetry in normally bilaterally symmetrical traits. We examined the relationship between FA of five body traits (ear height, length of three digits, and ankle circumference) and self-reported scores of physical and verbal aggression in a sample of 90 boys aged 10 to 15 years. The relationships between FA and scores of aggression (particularly physical aggression) were found to be negative; in other words, the most symmetrical boys showed highest aggression. One trait (ankle circumference) showed the characteristics of “ideal” FA—parametric mean of zero and a normal distribution. Mean asymmetries calculated from six repeated measures of ankle FA in 30 subjects taken over a period of five months showed strong negative associations with scores of physical aggression which were independent of age, height, and weight. It is argued that soft tissue “cyclical” FA (as opposed to “fixed” bony FA) is dependent on the secretion of hormones: for example, cortisol. Causal associations between behavioral traits such as aggresion and hormones will lead to similar correlations between FA and behavior. John Manning is a senior lecturer in the School of Biological Sciences at the University of Liverpool. His research interests are in symmetry as an indicator of good genes, disease resistance, fertility, and hormonal status in human and nonhuman animals. Daniel Wood has a B.Sc. in anatomy and biology from Liverpool University. At present he is a postgraduate student in the School of Biological Sciences. His research interests are in fluctuating asymmetry and its behavioral correlates in adolescent boys.     

Patterns of fluctuating asymmetry in flowers: Implications for sexual selection in plants

Characters in animals used in signalling and subjected to strong directional selection often demonstrate (i) an elevated level of fluctuating asymmetry (small random deviations from bilateral symmetry) and (ii) a negative relationship between the degree of individual fluctuating asymmetry and the size of a given character. We tested these two predictions in plants since flowers are subjected to strong directional selection and are involved in signalling to pollinators, whereas leaves are supposed not to be directly involved in signalling. The overall level of fluctuating asymmetry in a number of plant species with bilaterally or radially symmetric flowers was not generally higher in floral traits than in leaves. The level of fluctuating asymmetry in plants was sometimes significantly consistent within individuals. The absolute degree of individual fluctuating asymmetry in floral traits was generally negatively related to the size of the trait, while there was a positive relationship for leaves. The degree of individual fluctuating asymmetry in floral traits was marginally negatively related to the degree of individual fluctuating asymmetry in leaf traits. These patterns of fluctuating asymmetry in plants suggest that (i) the degree of asymmetry in flowers signals different aspects of quality than does the degree of asymmetry in leaves, and that (ii) fluctuating asymmetry in flowers often reflects the phenotypic quality of individual plants.     

SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

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 paradigm

A prominent paradigm in evolutionary biology over the last ten years has been the role of fluctuating asymmetry in sexual selection. Fluctuating asymmetry in bilaterally paired traits, and in particular sexual traits, has been proposed to be a reliable indicator of individual quality and the focus of selection through sexual competition and attractiveness. We surveyed the literature on fluctuating asymmetry and sexual selection and found a marked chronological decline in the proportion of studies supporting the paradigm through the 1990s. Our data are supported by an independent meta-analysis of the literature. The data conform with the notion of a scientific revolution in which the early phase of a paradigm change is characterized by a publication bias, a less critical approach to research, or both. The patterns we observe in the fluctuating asymmetry literature suggest caution in drawing general conclusions from meta-analyses conducted before revolutions have settled.     

Testosterone and Jamaican Fathers

This paper investigates relationships between men’s testosterone and family life in a sample of approximately 350 Jamaican fathers of children 18–24 months of age. The study recognizes the role of testosterone as a proximate mechanism coordinating and reflecting male life history allocations within specific family and cultural contexts. A sample of Jamaican fathers and/or father figures reported to an assessment center for an interview based on a standardized questionnaire and provided a saliva sample for measuring testosterone level. Outcomes measured include subject demographics such as age and relationship status as well as partnership quality and sexuality and paternal attitudes and behavior. The variation in these fathers’ relationship status (e.g., married co-residential fathers, fathers in new non-residential relationships) was not associated with men’s testosterone. Too few stepfathers participated to enable a direct test of the prediction that stepfathers would have higher testosterone than biological fathers, although fathers who reported living with partners’ (but not his own) children did not have higher testosterone than fathers not reporting residing with a non-biological child. Fathers’ relationship quality was negatively related to their testosterone. Measures of paternal attitudes and behavior were not related to fathers’ testosterone. Consistent with previous ethnography, this sample of Jamaican fathers exhibited variable life history profiles, including residential status. We discuss why fathers’ relationship quality was found to be negatively related to their testosterone level, but other predictions were not upheld.     

Influence of mother's heterozygosity on the variation of anthropometric traits in newborns]

The relationship between mother's heterozygosity as revealed by 14 polymorphic loci and variation of the set of anthropometric traits were studied in normal singleton newborns (174 boys, 127 girls). Statistically significant negative correlation between mother's heterozygosity and the index of fluctuating asymmetry was found in girls. Variances of four traits--body weight, body length, circumference of head and breast measured by first principal component, and mean number of minor deviations from development (stigma) are shown to be minimal in the children born by mothers with the average level of heterozygosity. The groups of newborns with different level of mother's heterozygosity are characterized by different combinations of first principal component, stigma, and the index of fluctuating asymmetry. It is concluded from the whole set of data that children born by mothers with the average level of heterozygosity have the highest level of viability.     

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.     

Fluctuating asymmetry in red junglefowl

Fluctuating asymmetry, the random deviation from perfect bilateral symmetry, has recently attracted considerable attention. Levels of asymmetry have been shown to correlate with measures of individual quality. We measured asymmetry in a variety of ornamental and non-ornamental traits in red junglefowl, Gallus gallus and examined the patterns of asymmetry among different traits within an individual. All ornamental traits had significantly higher levels of fluctuating asymmetry than did non-ornamental traits. However, inter-trait correlations of asymmetry were low for both ornamental and non-ornamental traits. We then correlated measures of asymmetry with several potential indicators of male quality, including comb size, body size, and body condition. We found little evidence that asymmetry in any measured trait reflected male quality. We measured asymmetry in ornamental traits at several stages of development and found no relationship between male condition and changes in asymmetry over time. Our results indicate that it is necessary to employ caution when choosing traits to be measured in studies of fluctuating asymmetry and that a relationship between asymmetry and individual quality cannot be assumed.     

Fluctuating asymmetry and fitness in Drosophila melanogaster

Models predict that developmental stability measured by fluctuating asymmetry should be positively correlated with fitness. Although such a correlation has often been suggested by indirect studies, there is still a lack of direct experimental evidence. In this note, I have measured the fluctuating asymmetry of sternopleural bristle counts in 32 lines of Drosophila melanogaster sharing the same genetic background but displaying all combinations of five visible mutations. Fluctuating asymmetry was heterogeneous among lines, suggesting a direct impact of the mutations on developmental stability. Two measures of fitness were made for each line: productivity (a combined measure of fecundity and egg‐to‐adult survivorship) and competitive male mating success. Fluctuating asymmetry was correlated with neither of these two components of fitness. This suggests that generalizations about fluctuating asymmetry must be taken with care.     

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.     

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.