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.     

Anatomic brain asymmetry in vervet monkeys

Asymmetry is a prominent feature of human brains with important functional consequences. Many asymmetric traits show population bias, but little is known about the genetic and environmental sources contributing to inter-individual variance. Anatomic asymmetry has been observed in Old World monkeys, but the evidence for the direction and extent of asymmetry is equivocal and only one study has estimated the genetic contributions to inter-individual variance. In this study we characterize a range of qualitative and quantitative asymmetry measures in structural brain MRIs acquired from an extended pedigree of Old World vervet monkeys (n = 357), and implement variance component methods to estimate the proportion of trait variance attributable to genetic and environmental sources. Four of six asymmetry measures show pedigree-level bias and one of the traits has a significant heritability estimate of about 30%. We also found that environmental variables more significantly influence the width of the right compared to the left prefrontal lobe.     

Response of fluctuating and directional asymmetry to selection on wing shape in Drosophila melanogaster

We tested whether directional selection on an index-based wing character in Drosophila melanogaster affected developmental stability and patterns of directional asymmetry. We selected for both an increase (up selection) and a decrease (down selection) of the index value on the left wing and compared patterns of fluctuating and directional asymmetry in the selection index and other wing traits across selection lines. Changes in fluctuating asymmetry across selection lines were predominantly small, but we observed a tendency for fluctuating asymmetry to decrease in the up-selected lines in both replicates. Because changes in fluctuating asymmetry depended on the direction of selection, and were not related to changes in trait size, these results fail to support existing hypotheses linking directional selection and developmental stability. Selection also produced a pattern of directional asymmetry that was similar in all selected lines whatever the direction of selection. This result may be interpreted as a release of genetic variance in directional asymmetry under selection.     

Heritabilities of Directional Asymmetry in the Fore- and Hindlimbs of Rabbit Fetuses

Directional asymmetry (DA), where at the population level symmetry differs from zero, has been reported in a wide range of traits and taxa, even for traits in which symmetry is expected to be the target of selection such as limbs or wings. In invertebrates, DA has been suggested to be non-adaptive. In vertebrates, there has been a wealth of research linking morphological asymmetry to behavioural lateralisation. On the other hand, the prenatal expression of DA and evidences for quantitative genetic variation for asymmetry may suggest it is not solely induced by differences in mechanic loading between sides. We estimate quantitative genetic variation of fetal limb asymmetry in a large dataset of rabbits. Our results showed a low but highly significant level of DA that is partially under genetic control for all traits, with forelimbs displaying higher levels of asymmetry. Genetic correlations were positive within limbs, but negative across bones of fore and hind limbs. Environmental correlations were positive for all, but smaller across fore and hind limbs. We discuss our results in light of the existence and maintenance of DA in locomotory traits.     

Inbreeding and developmental stability in three-spined sticklebacks (Gasterosteus aculeatus L.)

Fluctuating asymmetry, small non-directional departures from perfect symmetry in bilateral traits, results from the inability of individuals to buffer development against genetic and environmental perturbations. Fluctuating asymmetry is a widely used measure of developmental stability, and developmental stability has been hypothesised to be inversely related to heterozygosity. We compared male three-spined sticklebacks (Gasterosteus aculeatus L.) that had been inbred for one generation to outbred control males with respect to the asymmetry of a set of bilateral morphometric traits. Inbred fish developed significantly more asymmetric pectoral fins than their outbred counterparts, whereas neither the magnitude of asymmetry for pelvic spines nor for gill covers significantly responded to the treatment. Our results conform to a pattern of heterogeneity amongst traits in their tendency to develop asymmetrically in response to stress.     

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.     

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.     

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.     

Fluctuating asymmetry,body size and sexual selection in the dung fly Sepsis cynipsea — testing the good genes assumptions and predictions

In the dung fly Sepsis cynipsea large and more symmetric males have been shown to enjoy a mating advantage, but we still do not know which mechanism of sexual selection is responsible. Here we test several assumptions and predictions relating to the hypothesis that either trait is indicative of ‘good genes’. We tested for good genes by regressing fitness in good and bad environments (no and high larval competition, respectively) on the family mean for size or asymmetry as expressed in the good environment, separately for both sexes. Body size (hind tibia length or head width) was positively correlated with female fecundity, growth rate of both sexes and larval survivorship for males, but only in the good environment. The corresponding evidence for asymmetry is more equivocal. Mean standardised asymmetry was weakly associated with lower survivorship in the good environment, while growth rates and female fecundity were not. As predicted by sexual selection theory, fore tibia length showed greater asymmetry than other, presumably not sexually selected traits, and asymmetry in fore tibia length was greater for males than females. However, a negative correlation between trait size and asymmetry was only evident for male seta length but not for fore tibia length, fore femur length, or any composite measure of asymmetry. Most crucially, asymmetry was heritable for some female morphological traits (hind tibia length: h² = 0.15; fore femur length: h² = 0.16; mean of all measured traits: h² = 0.27), but not for any male trait. Also, asymmetry of the various traits measured was not correlated within males and only weakly so within females. The crucial assumption that asymmetry of sexually selected traits reflects overall, heritable developmental stability of an individual is thus only partly substantiated by our data. In contrast, large body size is heritable, associated with high fitness and consequently could be indicative of good genes. Fore leg asymmetry may influence male mating success by simply mechanically constraining his ability to hold on to the female.     

Assemblage and population-level consequences of forest fragmentation on bilateral asymmetry in tropical montane birds

Habitat fragmentation has the potential to influence the development and thus the phenotype of organisms. The asymmetry of bilateral traits may be indicative of the extent to which developmental stability is compromised by the stressful conditions underlying fragmentation. Using an assemblage- and population-level approach, we explored asymmetry differences in tarsus and outermost tail feathers of birds inhabiting fragmented landscapes in the tropical Andes of Colombia. More than 2500 individuals of 185 species were mist-netted at nine forest sites representing continuous forest (> 1000 ha), medium- (70–110 ha), and small-sized (8–20 ha) fragments. Feathers showed true fluctuating asymmetry (FA), whereas tarsus presented a mixture of FA and directional asymmetry. Overall, asymmetry was lowest in continuous forest, and highest in small and medium fragments. These patterns remained unchanged when directionality and differences in species composition, abundance, and foraging tactics were considered. The population-level analyses showed a general trend of increased asymmetry variation in fragments, yet the responses were not always in the same direction. Increased asymmetry may represent an outcome of processes that contribute to the persistence of species in changing environments, and to the generation of phenotypic innovation, which suggests individual adjustments of development to deal with stress. This calls into question the deliberated application of FA as a biomonitoring tool for conservation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 119–133.     

Developmental stability,sexual selection and speciation

Fluctuating asymmetry occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. Since both sides of a bilaterally symmetric trait are the result of the actions of a single genome, fluctuating asymmetry represents an epigenetic measure of the sensitivity of development to stress. Different morphological traits may show a direct relationship between their functional importance and their degree of developmental canalization. This may explain why some characters show high degrees of fluctuating asymmetry, and why these characters more often become exaggerated secondary sexual ornaments. The degree of fluctuating asymmetry is generally larger in small marginal populations living in novel environments, and this will particularly lead to relatively large degrees of asymmetry in the least developmentally canalized traits. More stringent selection against heterozygotes in marginal populations may further break down developmental stability and linkage groups which would lead to increased genetic variance. Females may prefer to mate with males having large, but relatively symmetric morphological characters, because it is more difficult to make large traits (a good genes argument), a large trait is more easily perceived (a sensory bias preference), and because symmetry signals ability to cope with stress (a good genes argument). The low degree of developmental stability and the large amount of genetic variance in secondary sexual characters in small, marginal populations could set the scene for rapid development of divergence and speciation in marginal populations.     

Developmental instability in Brassica campestris (Cruciferae): fluctuating asymmetry of foliar and floral traits

The degree of fluctuating asymmetry of bilateral traits provides a measure of developmental instability, which can be influenced by genetic as well as environmental stress. We studied genetic variation between and within two populations of the mustard Brassica campestris for asymmetry of foliar (cotyledon width) and floral (petal length and width) traits as well as for phenological (germination and flowering) and performance (biomass and flowering) traits. The two populations differed in mean expression of most traits, including asymmetry. However, within-population estimates of genetic variability tended to be lower for asymmetry than other traits. Asymmetry was greater in the population that had lower biomass accumulation and flower production, which supports the idea that population-level asymmetry may be indicative of population-level performance. However, within each population, evidence that performance was negatively correlated with asymmetry was equivocal. Within populations there was little or no concordance among estimates of asymmetry based on different structures, i.e., plants that had highly asymmetrical cotyledons did not tend to have highly asymmetrical petals. The lack of a general buffering capacity at the individual level may be explained by developmental processes (e.g., action of different genes or morphogens) as well as evolutionary processes (e.g., selection on asymmetry of different traits).     

Testes asymmetry,condition and sexual selection in birds: an experimental test

The functional significance of the marked directional asymmetry in testes size observed in many bird species is obscure. Møller suggested that (i) the smaller of the two testes serves a compensatory role and increases in size (and hence reduces asymmetry) when the larger one is defective in some way, and (ii) as a consequence, the degree of directional asymmetry in testes size reflects male quality and covaries positively with the expression of secondary sexual traits. We conducted an experimental test of these two hypotheses in the zebra finch, Taeniopygia guttata. Neither hypothesis was supported. First, there was no significant relationship between the size of the left testis and relative testes asymmetry. Second, we obtained no support for the hypothesis that the degree of directional asymmetry in testes mass covaried with condition. On the contrary, directional asymmetry in testes mass was significantly greater in birds whose condition was experimentally reduced, compared with control birds. Moreover, we found no significant relationships between testes asymmetry and secondary sexual traits. We conclude that directional asymmetry in testes size does not reflect male condition in the zebra finch.     

Fingerprint quantitative variation and asymmetry in Brazilian Whites and Blacks

A total of 432 Whites and 654 Blacks were studied in relation to the index of pattern intensity and a smaller number (296 Whites; 496 Blacks) for total ridge count. There does not exist a clear gradient in either sex when the Blacks are subdivided into three phenotypic categories according to the amount of African ancestry they seem to possess. The standard deviations and coefficients of variation are strikingly similar in all groups studied. Racial differences in the averages of these two quantitative measures and in the asymmetry of total ridge count are small and nonsignificant. Fingerprints are very sensitive indicators of disturbances in the intrauterine development, and their asymmetry has a significant hereditary component. Since there is no increased or decreased asymmetry when we consider subgroups with various amounts of racial admixture, these and other results suggest that genetic adaptation in humans is a process involving mainly the whole species, and not narrow racial specializations.     

Correcting for allometry in studies of fluctuating asymmetry and quality within samples

Fluctuating asymmetry (FA) refers to deviations from perfect symmetry in a bilateral character and is believed to reflect an organism''s quality. However, allometric relations between asymmetry and trait size may confound FA–quality relations. Larger traits may have more ''opportunity'' to become asymmetric. Thus, researchers suggest that if allometric relations between FA and size are found, some correction for size is needed (typically FA scaled by size). I used a simulation model to examine potential consequences of allometry for detection of FA–quality relations within samples and examined efficacy of rules currently used to correct for size. Consequences of allometry can be severe, causing up to 77% type I errors or 98% power loss when true FA–quality relations exist (when FA–size relations are isometric), depending upon the strength and direction of size–quality relations. I illustrate why current rules to correct for size are inefficient and suggest alternative rules. Problems of allometry that require further attention are discussed.     

Polymorphism and multiple correlated characters: Do flatfish asymmetry morphs also differ in swimming performance and metabolic rate?

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Patterns of fluctuating asymmetry in sexual ornaments predict female choice

Extravagant secondary sexual characters are assumed to have arisen and be maintained by sexual selection. While traits like horns, antlers and spurs can be ascribed to intrasexual competition, other traits such as extravagant feather ornaments, displays and pheromones have to be ascribed to mate choice. A number of studies have tested whether females exert selection on the size of male ornaments, but only some of these have recorded female preferences for the most extravagantly ornamented males. Here I demonstrate that female choice can be directly predicted from the relationship between the degree of fluctuating asymmetry and the size of a secondary sexual character. Fluctuating asymmetry is an epigenetic measure of the ability of individuals to cope with stress, and it occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. There is a negative relationship between the degree of fluctuating asymmetry and the absolute size of an ornament in those bird species with a female preference for the largest male sex trait, while there is a flat or U-shaped relationship among species without a female preference. These results suggest that females prefer exaggerated secondary sexual characters if they reliably demonstrate the ability of males to cope with genetic and environmental stress. Some species may demonstrate a flat or U-shaped relationship between the degree of fluctuating asymmetry and the absolute size of an ornament because (i) the genetic variance in viability signalled by the secondary sex trait has been depleted; (ii) the secondary sex trait is not particularly costly and therefore does not demonstrate condition dependence; or because (iii) the sex traits can be considered arbitrary traits rather than characters reflecting good genes.     

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.     

Heterozygosity and asymmetry: Ectodysplasin as a form of genetic stress in marine threespine stickleback

Genome‐wide heterozygosity has long been hypothesized to play a role in buffering organisms against developmental perturbations, potentially resulting in increased symmetry. If true, this could in part explain the maintenance of standing genetic variation in wild populations. Marine threespine sticklebacks (Gasterosteus aculeatus) were sampled across their eastern Pacific coastal distribution from Alaska to California and variations in asymmetry for both structural and nonstructural armor traits (lateral plates) were assessed. Structural plates consistently showed less asymmetry than nonstructural plates, but standardized measures of heterozygosity were not correlated with the extent of asymmetry expressed by a fish. Fish that were heterozygous for the major‐effect gene controlling lateral plate variation (Ectodysplasin) had higher occurrences of asymmetry, even when the individuals were phenotypically fully plated. Collectively, this suggests that heterozygosity at a major‐effect locus can have a greater impact on asymmetry than heterozygosity sampled across the genome.     

Genetic correlations between sides and heritability of asymmetry for nonmetric traits in rhesus macaques on Cayo Santiago

The use of nonmetric traits for estimation of biological distance is a long-standing practice in biological anthropology. Nonmetric traits can be scored using either the individual or the side of the individual as the unit of measure. If sides of the individual are genetically correlated the use of sides would produce redundant genetic information. For this reason, Korey (Am. J. Phys. Anthropol. 53:19-23, 1980) argues for the use of individuals as the unit of measure for nonmetric traits. Ossenberg (Am. J. Phys, Anthropol. 54:471-479, 1981), however, argues that bilateral occurrence of nonmetric traits indicates greater genetic liability for the trait and that therefore the sides are the more biologically correct unit of measure. Genetic correlations for 13 cranial nonmetric traits are estimated for a sample of rhesus macaque skeletons from Cayo Santiago. In addition, heritability of asymmetry is estimated for these 13 traits as a test of Ossenberg's contention that asymmetry is genetically influenced. Significant genetic correlations between sides support Korey's contention that nonmetric traits should be scored by individual. Only two asymmetry heritabilities were significantly different from zero, providing no significant support for Ossenberg's contention that asymmetry is genetically determined. Our results support the theory that asymmetry represents a measure of the ability of an organism to buffer stresses. Therefore, a measure of the heritability of asymmetry is a measure of the heritability of the ability to buffer stresses. This ability does not appear to be heritable in this sample.