Variation In Fluctuating Asymmetry Research Articles

Relationship between flight muscle dimorphism and wing morphometry in Triatoma infestans (Klug, 1834) (Hemiptera, Reduviidae, Triatominae)

Triatoma infestans, the primary vector of Chagas disease in southern South America, is reported as a dimorphic species for flight muscle development and presents a marked phenotypic plasticity in traits related to flight. In triatomines, studying the relationship between flight muscle development and wing variation is important, since it involves flight capacity and dispersion. Therefore, we examined the association between flight muscle dimorphism and morphological wing variation in T. infestans individuals. We hypothesized that if the underdeveloped flight muscle were associated with a decrease in wing developmental stability, canalization and morphological aspect of wings, and with a loss of wing modularity, then wing variation would be associated with flight muscle dimorphism. Wing size and shape were characterized using geometric morphometrics. Patterns of canalization, developmental instability, and modularity were inferred from fluctuating asymmetry analysis of wing shape. Morphological aspects of wings were estimated using maximum wing length and wing area. The results showed that there are differences in wing shape but not in wing size among groups. Canalization and developmental stability were positively associated in individuals with muscle development and negatively associated in individuals with underdeveloped flight muscles. The covariance structure of variation among individuals was different between groups but not for fluctuating asymmetry, both for females and males. Modularity hypothesis related to the wing subdivision into two parts, a stiff part and a membranous one, showed significant association for the variation among individuals and fluctuating asymmetry variation in females with flight muscles. Wing aspect showed that females with flight muscles exhibited more elongated and thinner wings than those without flight muscles. The results obtained in this study reveal an association between flight muscle development and forewing variation in T. infestans. Females with flight muscles have certain differential characteristics in their wings that would be related to an enhanced flight performance.

Differential Impact of Forest Fragmentation on Fluctuating Asymmetry in South Amazonian Small Mammals

(1) Background: Southern Amazonia encompasses some of the most human-impacted and deforested regions of South America, resulting in a hyper-fragmented landscape. In this context, by using a geometric morphometrics approach, we aimed to examine the effect of forest fragmentation on developmental instability (DI) of the mandible, assessed by variation of fluctuating asymmetry (FA), in four neotropical small mammal species inhabiting the municipality of Alta Floresta (Brazil). (2) Methods: The impact of fragment area, fragment shape, isolation, and edge length on DI were assessed by measuring variation in mandibular FA in the long-tailed spiny rat (Proechimys longicaudatus), the hairy-tailed bolo mouse (Necromys lasiurus), the woolly mouse opossum (Marmosa demerarae), and the Amazonian red-sided opossum (Monodelphis glirina). Mandibles from a total of 304 specimens originating from different-sized fragments (ranging from 5 to 900 ha) were used. Twelve homologous landmarks were digitized in photographs of the mesial view of each hemi-mandible. (3) Results: The two largest species, P. longicaudatus and M. demerarae, exhibited significantly higher levels of FA in mandibular shape in small fragments (5–26 ha) in comparison to large ones (189–900 ha). Edge length negatively impacted M. demerarae, the only arboreal species, reinforcing its strongest dependence on core forest habitats. (4) Conclusions: For small mammal communities, we propose that fragments >~200 ha should be the focus of conservation efforts, as both resilient and more sensitive species would benefit from their more preserved biotic and abiotic conditions. Conversely, fragments <~25 ha seem to lead to a significant increase in stress during developmental stages.

Craniofacial fluctuating asymmetry in gorillas, chimpanzees, and macaques.

Craniofacial fluctuating asymmetry (FA) refers to the random deviations from symmetry exhibited across the craniofacial complex and can be used as a measure of developmental instability for organisms with bilateral symmetry. This article addresses the lack of data on craniofacial FA in nonhuman primates by analyzing FA magnitude and variation in chimpanzees, gorillas, and macaques. We offer a preliminary investigation into how FA, as a proxy for developmental instability, varies within and among nonhuman primates. We generated 3D surface models of 121 crania from Pan troglodytes troglodytes, Gorilla gorilla gorilla, and Macaca fascicularis fascicularis. Using geometric morphometric techniques, the magnitude of observed FA was calculated and compared for each individual, sex, and taxon, along with the variation of FA across cranial regions and for each bilateral landmark. Gorillas and macaques exhibited higher and more similar magnitudes of FA to each other than either taxon did to chimpanzees; variation in magnitude of FA followed this same trend. No significant differences were detected between sexes using pooled data across species, but sex did influence FA magnitude within taxa in gorillas. Further, variation in FA variance across cranial regions and by landmark was not distributed in any particular pattern. Possible environmentally induced causes for these patterns of FA magnitude include differences in growth rate and physiological stress experienced during life. Developmental stability may be greatest in chimpanzees in this sample. Additionally, these results point to appropriate landmarks for future FA analyses and may help suggest more urgent candidate taxa for conservation efforts.

Variation in Leaf Size and Fluctuating Asymmetry of Mountain Birch (Betula pubescens var. pumila) in Space and Time: Implications for Global Change Research

Experimental, latitudinal, and historical approaches have been used to explore and/or predict the effects of global change on biota, and each approach has its own advantages and disadvantages. The weaknesses of these individual approaches can, potentially, be avoided by applying them simultaneously, but this is rarely done in global change research. Here, we explored the temporal and spatial variations in the leaf size and fluctuating asymmetry (FA) of mountain birch (Betula pubescens var. pumila) in the Murmansk region of Russia, with the aim of verifying the predictions derived from the responses of these traits to experimental manipulations of abiotic drivers of global change. The examination of herbarium specimens revealed that leaf length increased during the 20th century, whereas the FA in the number of leaf teeth decreased, presumably reflecting an increase in the carbon and nitrogen availability to plants in that century. Along a northward latitudinal gradient, leaf length decreased whereas FA increased, presumably due to the poleward decreases in air temperature. The study site, collection year, and latitude explained a larger part of the leaf length variation in mountain birch relative to the variation in FA. Leaf length is likely a better indicator than FA in studies addressing global environmental change impacts on plant performance.

Mouse Skull Mean Shape and Shape Robustness Rely on Different Genetic Architectures and Different Loci.

The genetic architecture of skull shape has been extensively studied in mice and the results suggest a highly polygenic and additive basis. In contrast few studies have explored the genetic basis of the skull variability. Canalization and developmental stability are the two components of phenotypic robustness. They have been proposed to be emergent properties of the genetic networks underlying the development of the trait itself, but this hypothesis has been rarely tested empirically. Here we use outbred mice to investigate the genetic architecture of canalization of the skull shape by implementing a genome-wide marginal epistatic test on 3D geometric morphometric data. The same data set had been used previously to explore the genetic architecture of the skull mean shape and its developmental stability. Here, we address two questions: (1) Are changes in mean shape and changes in shape variance associated with the same genomic regions? and (2) Do canalization and developmental stability rely on the same loci and genetic architecture and do they involve the same patterns of shape variation? We found that unlike skull mean shape, among-individual shape variance and fluctuating asymmetry (FA) show a total lack of additive effects. They are both associated with complex networks of epistatic interactions involving many genes (protein-coding and regulatory elements). Remarkably, none of the genomic loci affecting mean shape contribute these networks despite their enrichment for genes involved in craniofacial variation and diseases. We also found that the patterns of shape FA and individual variation are largely similar and rely on similar multilocus epistatic genetic networks, suggesting that the processes channeling variation within and among individuals are largely common. However, the loci involved in these two networks are completely different. This in turn underlines the difference in the origin of the variation at these two levels, and points at buffering processes that may be specific to each level.

Does fluctuating asymmetry of hind legs impose costs on escape speed in house crickets (Acheta domesticus)?

Fluctuating asymmetry (FA) is often thought to be an indicator of developmental stability—an individual’s ability to resist environmental and genetic stress during development—and thus demonstrates phenotypic quality. Research on the influence of FA on locomotion has often found that high FA in legs and wings impedes locomotor performance. Crickets rely on their six limbs to flee from predators and parasitoids. Hind legs are of particular importance during escape as they contribute to running speed. FA research overwhelmingly focuses on its impact on sexual selection, with little on locomotion and only one study of the impact of FA on invertebrate locomotion. Here, we examined the effect of FA in hind legs on the escape speed of house crickets (Acheta domesticus) and the locomotor costs of hind limb autotomy. Unexpectedly, our findings indicate that FA of hind legs have no influence on the escape speed of either male or female A. domesticus. This is inconsistent with most research conducted on FA and vertebrate locomotion that indicates FA negatively impacts locomotion, but is consistent with the only research examining FA and invertebrate locomotion. Our other findings were more congruent with the literature on other Orthoptera, as body size was found to have an influence on the escape speeds of intact females and those that lost two hind limbs. Whilst our results indicate that FA did not influence locomotion, this may not be the case for other invertebrate taxa where variation in FA may have an important role in natural selection.

Do defoliating insects distinguish between symmetric and asymmetric leaves within a plant?

1. Plants represent a highly heterogeneous resource for herbivores. One dimension of this heterogeneity is reflected by the within‐plant variation in the leaf fluctuating asymmetry (FA), i.e. in the magnitude of the random deviations from the symmetry in leaf shape.2. This study is the first to test experimentally the hypothesis that variation in the quality of individual leaves for defoliating insects (11 species) within a plant (seven tree and shrub species) is associated with the FA of these leaves.3. It was demonstrated that specialist defoliators generally distinguish between nearly symmetric (low FA) and highly asymmetric (high FA) leaves, but do not distinguish between discs cut from these leaves. Low‐FA leaves of Salix caprea, Salix myrsinifolia and Populus tremula were of better quality for insects than high‐FA leaves, as demonstrated by both preference tests and performance trials. By contrast, high‐FA leaves of Betula pubescens were of better quality for insects than low‐FA leaves, whereas insects feeding on Alnus incana showed similar responses to high‐ and low‐FA leaves.4. It is concluded that insect herbivores can distinguish between leaves with high and low FA, and that FA may be associated with the quality of an individual leaf for insects, although the direction and strength of the effect of leaf FA on insect preference and performance vary among study systems. The ecological significance of substantial within‐plant variation in leaf FA remains to be explored.

Local Insect Damage Reduces Fluctuating Asymmetry in Next-year’s Leaves of Downy Birch

Insect herbivory imposes stress on host plants. This stress may cause an increase in leaf fluctuating asymmetry (FA), which is defined as the magnitude of the random deviations from a symmetrical leaf shape. We tested the hypothesis that differences in leaf FA among individual shoots of downy birch, Betula pubescens, are at least partly explained by local damage caused by insects in the previous year. Unexpectedly, we found that in the year following the damage imposed by miners, leafrollers and defoliators, damaged birch shoots produced leaves with lower FAs compared to shoots from the same tree that had not been damaged by insects. This effect was consistent among the different groups of insects investigated, but intra-species comparisons showed that statistical significance was reached only in shoots that had been damaged by the birch leaf roller, Deporaus betulae. The detected decrease in leaf FA in the year following the damage agrees with the increases in shoot performance and in antiherbivore defence. The present results indicate that within-plant variation in leaf FA may have its origin in previous-year damage by insects, and that FA may influence the current-year’s distribution of herbivory.

Genetic structure of phenotypic robustness in the collaborative cross mouse diallel panel.

Developmental stability and canalization describe the ability of developmental systems to minimize phenotypic variation in the face of stochastic micro-environmental effects, genetic variation and environmental influences. Canalization is the ability to minimize the effects of genetic or environmental effects, whereas developmental stability is the ability to minimize the effects of micro-environmental effects within individuals. Despite much attention, the mechanisms that underlie these two components of phenotypic robustness remain unknown. We investigated the genetic structure of phenotypic robustness in the collaborative cross (CC) mouse reference population. We analysed the magnitude of fluctuating asymmetry (FA) and among-individual variation of cranial shape in reciprocal crosses among the eight parental strains, using geometric morphometrics and a diallel analysis based on a Bayesian approach. Significant differences among genotypes were found for both measures, although they were poorly correlated at the level of individuals. An overall positive effect of inbreeding was found for both components of variation. The strain CAST/EiJ exerted a positive additive effect on FA and, to a lesser extent, among-individual variance. Sex- and other strain-specific effects were not significant. Neither FA nor among-individual variation was associated with phenotypic extremeness. Our results support the existence of genetic variation for both developmental stability and canalization. This finding is important because robustness is a key feature of developmental systems. Our finding that robustness is not related to phenotypic extremeness is consistent with theoretical work that suggests that its relationship to stabilizing selection is not straightforward.

How reproducible are the measurements of leaf fluctuating asymmetry?

Fluctuating asymmetry (FA) represents small, non-directional deviations from perfect symmetry in morphological characters. FA is generally assumed to increase in response to stress; therefore, FA is frequently used in ecological studies as an index of environmental or genetic stress experienced by an organism. The values of FA are usually small, and therefore the reliable detection of FA requires precise measurements. The reproducibility of fluctuating asymmetry (FA) was explored by comparing the results of measurements of scanned images of 100 leaves of downy birch (Betula pubescens) conducted by 31 volunteer scientists experienced in studying plant FA. The median values of FA varied significantly among the participants, from 0.000 to 0.074, and the coefficients of variation in FA for individual leaves ranged from 25% to 179%. The overall reproducibility of the results among the participants was rather low (0.074). Variation in instruments and methods used by the participants had little effect on the reported FA values, but the reproducibility of the measurements increased by 30% following exclusion of data provided by seven participants who had modified the suggested protocol for leaf measurements. The scientists working with plant FA are advised to pay utmost attention to adequate and detailed description of their data acquisition protocols in their forthcoming publications, because all characteristics of instruments and methods need to be controlled to increase the quality and reproducibility of the data. Whenever possible, the images of all measured objects and the results of primary measurements should be published as electronic appendices to scientific papers.

Lack of response to artificial selection on developmental stability of partial wing shape components in Drosophila melanogaster.

Developmental stability, the ability of organisms to buffer their developmental processes against developmental noise is often evaluated with fluctuating asymmetry (FA). Natural genetic variation in FA has been investigated using Drosophila wings as a model system and the recent estimation of the heritability of wing shape FA was as large as 20%. Because natural genetic variation in wing shape FA was found to localize in a partial component of the wings, heritable variation in specific parts of the wings might be responsible for FA estimation based on the whole wing shape. In this study, we quantified the shape of three partial components of the wings, and estimated the heritability of the wing shape FA based on artificial selections. As a result, FA values for the partial wing shape components did not respond to artificial selections and the heritability scores estimated were very small. These results indicate that natural additive genetic variation in FA of partial wing components was very small compared with that in a complex wing trait.

Fluctuating asymmetry and Echinospartum horridum fitness components

In the last two decades, fluctuating asymmetry (FA) has been used commonly as an indicator of the stress experienced by individual organisms during development, although the technique has not been without controversy. In general, estimates of FA in populations have been positively correlated with measures of stress but, in the few studies in which it has been investigated, the correlations between estimates of FA and stress in individuals have been inconsistent. This study compared the FA in the opposite spines of stems and flowers to other bio-indicators of physiological stress in plants, such as annual crown growth ratio, annual flower production, annual crown death ratio, and branch fractal dimension. The position of an individual Echinospartum horridum within a patch influenced the stress conditions, and influenced its colonization strategy. Plants in the interior of patches are under high intraspecific competition due to negative density feedback (low flower production and branch biomass, and high annual crown death ratio), showed larger flower FA and spines FA repeatability (within-individual variation in FA) than plants at the periphery. Plant growth rates and the FA of spines were negatively correlated, but flower asymmetry and flower production was not significantly correlated, which reflects the capacity of E. horridum to adjust flower production to its developing conditions. We concluded that because organisms respond differently to different conditions, enhanced by the plasticity of plants in the development of morphological structures in response to changes in the environment, it is advisable to use multiple parameters to assess physiological stress in plants.

Insect Herbivores ofCoccoloba cereiferaDo Not Select Asymmetric Plants

Fluctuating asymmetry (FA) represents small, random variations from the symmetric bilateral pattern, and it is a frequently used measurement of plant and animal stresses. Some studies have shown a relationship between FA and herbivory, suggesting that FA might be a reliable indicator of plant quality and plant susceptibility to herbivore attack. In this study, we investigated the relationship between leaf FA of Coccoloba cereifera Schwacke (Polygonaceae) and the pattern of attack by the scale insect Abgrallaspis cyanophylly Signoret (Hemiptera: Diaspididae). Thirty individuals of C. cereifera were marked and followed during 2007 in southeastern Brazil, and all leaves of three shoots per plant were sampled, numbered, and categorized as young, mature, or senescent. We measured right and left widths of all leaves after digitizing the images to determine levels of FA for each leaf sampled per plant. Individuals of A. cyanophylli were counted using a stereoscope in the laboratory to determine infestation levels. Leaves of C. cereifera exhibited true patterns of FA, as shown by the normal distribution of asymmetry values. We observed high variation in attack rates by A. cyanophylly among C. cereifera individuals, with 43.3% of plants with no insects, whereas others exhibited up to 23,000 insects. No statistically significant relationship was observed between variation in FA and variation in A. cyanophylli among plants. These results suggest that FA cannot be used as an indicator of plant quality and susceptibility to herbivore attack but suggest that herbivores themselves might act as plant stressors.

Small Variations over Large Scales: Fluctuating Asymmetry over the Range of Two Oak Species

Fluctuating asymmetry (FA) represents small, random variation from symmetry, and it has been used as an indicator of plant stress at relatively local scales. In this study, variation in FA was examined at the landscape scale across the whole distributional range of the two oak species Quercus myrtifolia and Quercus chapmanii in Florida. Oak leaf morphology was evaluated in 40 sites covering ∼170,000 km2. Plants growing on coastal/edge sites exhibited significantly higher levels of FA than did plants inhabiting inland/center sites. For myrtle oaks, levels of FA were also spatially structured, and sites were positively spatially autocorrelated at small distances, indicating that sites that were closer exhibited similar levels of environmental stress and FA. Our results have shown that FA, a reliable measurement of plant stress at local scales, can also be used as a biological tool for monitoring the quality of the environment at larger spatial scales, increasing its usefulness in studies across the full ran...

Effects of small Hsp genes on developmental stability and microenvironmental canalization.

BackgroundProgression of development has to be insulated from the damaging impacts of environmental and genetic perturbations to produce highly predictable phenotypes. Molecular chaperones, such as the heat shock proteins (HSPs), are known to buffer various environmental stresses, and are deeply involved in protein homeostasis. These characteristics of HSPs imply that they might affect developmental buffering and canalization.ResultsWe examined the role of nine Hsp genes using the GAL4/UAS-RNAi system on phenotypic variation of various morphological traits in Drosophila melanogaster. The stability of bristle number, wing size and wing shape was characterized through fluctuating asymmetry (FA) and the coefficient of variation (CV), or among-individual variation. Progeny of the GAL4/Hsp-RNAi crosses tended to have reduced trait means for both wing size and wing shape. Transcriptional knockdown of Hsp67Bc and Hsp22 significantly increased FA of bristle number, while knockdown of Hsp67Ba significantly increased FA and among-individual variation of wing shape but only in males. Suppression of Hsp67Bb expression significantly increased among-individual variation of bristle number. The knockdown of gene expression was confirmed for Hsp67Ba, Hsp67Bc, Hsp22, and Hsp67Bb. Correlation between FA and CV or among-individual variation of each trait is weak and not significant except for the case of male wing shape.ConclusionFour small Hsp genes (Hsp22, Hsp67Ba, Hsp67Bb and Hsp67Bc) showed involvement in the processes of morphogenesis and developmental stability. Due to possible different functions in terms of developmental buffering of these small Hsps, phenotypic stability of an organism is probably maintained by multiple mechanisms triggered by different environmental and genetic stresses on different traits. This novel finding may lead to a better understanding of non-Hsp90 molecular mechanisms controlling variability in morphological traits.

Fluctuating Asymmetry in Common Tern (Sterna hirundo) Chicks Varies with Hatching Order and Clutch Size

ABSTRACT. Fluctuating asymmetry (FA), small random deviation from bilateral symmetry, often increases with stress during development. Common Terns (Sterna hirundo) typically lay two to three eggs that hatch asynchronously. I predicted that C-chicks (last of three) should have greater FA than A- and B-chicks at hatching and that FA should be higher in chicks from smaller clutches, because of differences in parental quality. Tarsus length of newly hatched chicks was measured across three years, and middle toe length was measured in one year. Sample sizes exceeded 100 chicks in two of three years. Variation in tarsus FA with hatching order and clutch size was statistically significant in one year (P < 0.01) and nearly so in another (P < 0.10). No significant differences were present for toe FA. A-chicks from three-egg clutches appeared to have the lowest tarsus FA among categories of chicks in both years, and in one year they were significantly more symmetrical than B- and C-chicks from three-egg clutches. A...

Evolutionary history shapes the association between developmental instability and population‐level genetic variation in three‐spined sticklebacks

Developmental instability (DI) is the sensitivity of a developing trait to random noise and can be measured by degrees of directionally random asymmetry [fluctuating asymmetry (FA)]. FA has been shown to increase with loss of genetic variation and inbreeding as measures of genetic stress, but associations vary among studies. Directional selection and evolutionary change of traits have been hypothesized to increase the average levels of FA of these traits and to increase the association strength between FA and population-level genetic variation. We test these two hypotheses in three-spined stickleback (Gasterosteus aculeatus L.) populations that recently colonized the freshwater habitat. Some traits, like lateral bone plates, length of the pelvic spine, frontal gill rakers and eye size, evolved in response to selection regimes during colonization. Other traits, like distal gill rakers and number of pelvic fin rays, did not show such phenotypic shifts. Contrary to a priori predictions, average FA did not systematically increase in traits that were under presumed directional selection, and the increases observed in a few traits were likely to be attributable to other factors. However, traits under directional selection did show a weak but significantly stronger negative association between FA and selectively neutral genetic variation at the population level compared with the traits that did not show an evolutionary change during colonization. These results support our second prediction, providing evidence that selection history can shape associations between DI and population-level genetic variation at neutral markers, which potentially reflect genetic stress. We argue that this might explain at least some of the observed heterogeneities in the patterns of asymmetry.

Fitness, developmental instability, and the ontogeny of fluctuating asymmetry in Daphnia magna

Fluctuating asymmetry (FA) is the most commonly used measure of developmental instability. The relation between FA and individual fitness remains controversial, partly due to limited knowledge on the mechanisms behind variation in FA. To address this, we investigated the associations between FA, growth and reproduction as well as the ontogeny of FA in a clonal population of Daphnia magna. FA was not correlated with growth and reproduction, either at the between- or the within-individual level, in a high (N = 48 individuals) or in a low (N = 52 individuals) food-quantity regime. There were therefore no indications of functional effects of FA or of phenotypic trade-offs between developmental stability, growth and reproduction. Individual asymmetries varied randomly in sign and magnitude between subsequent molts (N = 19 individuals, 9–11 instars), but the levels of FA were generally lowest at intermediate ages. No feedback between right and left sides was detected. This suggests that FA only reflects the most recent growth history, that developmental instability may increase in old age, and that FA depends on processes operating on each side of the body independently. The results also suggest that FA differences within and among individual Daphnia are largely random, with limited biological significance. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 179–192.

The Genetics and Evolution of Fluctuating Asymmetry

▪ Abstract Variation in the subtle differences between right and left sides of bilateral characters, or fluctuating asymmetry (FA), has long been considered to be primarily environmental in origin, and this has promoted its use as a measure of developmental instability (DI) in populations. There is little evidence for specific genes that govern FA per se. Numerous studies show that FA levels in various characters are influenced by dominance and especially epistatic interactions among genes. An epistatic genetic basis for FA may complicate its primary use in comparisons of DI levels in outbred or wild populations subjected or not subjected to various environmental stressors. Although the heritability of FA typically is very low or zero, epistasis can generate additive genetic variation for FA that may allow it to evolve especially in populations subjected to bottlenecks, hybridizations, or periods of rapid environmental changes caused by various stresses.

Lack of response to selection for lower fluctuating asymmetry of mutant eyespots in the butterfly Bicyclus anynana.

Fluctuating asymmetry (FA) is claimed both to provide a means of evaluating developmental stability, and to reflect an individual's quality or the stress experienced during development. FA refers to the nondirectional variation between left and right sides, whereas directional asymmetry (DA) refers to a significant directional variation between the sides. We studied four eyespots on the dorsal forewing of the tropical butterfly, Bicyclus anynana. Two of the eyespots were specified by a mutant allele, Spotty, that was fixed in the stock. These eyespots showed higher FA than the two flanking, wild-type eyespots, although they are all formed by the same developmental pathway. We applied artificial selection for lower FA of the novel eyespots in an attempt to increase their developmental stability. There was significant variation present in individual FA in our study. However, this did not change as a result of the artificial selection. Most of the variation in FA can be accounted for by individual differences in developmental stability rather than by the applied selection or by environmental variation. Thus, it was not possible to produce any increased developmental stability of the novel eyespots by selecting for low FA. The estimates of realized heritability for both FA and DA of each eyespot were not significantly different from zero. The results suggest that FA provides little, if any, potential for exploring the mechanistic basis of developmental stability.