Fluctuating asymmetry and stress in a medieval Nubian population

Fluctuating asymmetry is commonly used as a bioindicator of developmental stress. This study addresses asymmetry under nutritional/systemic stress in the human craniofacial skeleton and its utility as an indicator of developmental instability. Crania from the diachronic Christian cemeteries at Kulubnarti (Sudanese Nubia) were chosen as a model for nutrition/systemic stress. Previous studies indicate that individuals from the Early Christian cemetery were subjected to greater developmental stress when compared with individuals from the Late Christian cemetery. Therefore, crania from the Early Christian cemetery should display a greater magnitude of fluctuating asymmetry than crania from the Late Christian cemetery. Thirty adult crania of comparable age and sex were selected from each population. Landmark coordinates were digitized in two separate trials and averaged to minimize error. Euclidean distance matrix analysis (EDMA) was used to measure and compare the magnitude of fluctuating asymmetry in each sample. Results indicate that crania from the Early Christian cemetery display greater amounts of fluctuating asymmetry than those from the Late Christian cemetery, as predicted. The degree of fluctuating asymmetry for each linear distance is highly correlated between the cemeteries, suggesting that all humans may share common patterns of fluctuating asymmetry in the skull. In contrast, there is little correlation between magnitude of fluctuating asymmetry and length of linear distance, between-subject variability, or measurement error. These results support the hypothesis that poor nutrition/systemic stress increases developmental instability in the human skull and that increased fluctuating asymmetry constitutes morphological evidence of this stress.     

Fluctuating asymmetry as a possible measure of developmental homeostasis in humans: a review

We investigate three hypotheses related to fluctuating asymmetry (FA) of bilateral morphologic traits in humans: (1) the magnitude of FA in individual suffering from different levels of morbidity is significantly elevated compared with FA in healthy control subjects, (2) FA is negatively correlated with an individual's heterozygosity, and (3) phenotypic variance of FA may have a significant genetic component (or at least a family resemblance). Our experimental data and the literature support the first hypothesis and indicate that individuals who suffer from chromosomal or polygenic morbidity and from anomalies or conditions of development with still unknown genetic components demonstrate an elevated FA of various structures. The literature regarding the second hypothesis is sparse but is generally in agreement with it, although some exceptions exist. A study of correlations of phenotypic scores of FA between family members of nuclear families in two independent samples has shown that FA variance in individual traits probably does not have any significant genetic component. However, phenotypic variance of the mean estimate of FA over 8 traits showed significant additive and nonadditive (dominance) genetic components, each about 0.30.     

Fluctuating asymmetry as an indicator of stress: Implications for conservation biology

Extinction can be attributed broadly to environmental or genetic stress. The ability to detect such stresses before they seriously affect a population can enhance the effectiveness of conservation programs. Recent studies have shown that within-individual morphological variability may provide a valuable early indicator of environmental and genetic stress.     

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.     

Fluctuating asymmetry and sexual selection

Behavioral ecologists are being attracted to the study of within-individual morphological variability, manifested in random deviations from bilateral symmetry, as a means of ascertaining the stress susceptibility of developmental regulatory mechanisms. Several early successes Indicate that incorporating measures of symmetry into sexual-selection studies may help link individual sexual success to a basic component of viability - developmental stability.     

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 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.     

Fluctuating asymmetry and immune function in a field cricket

Recently, fluctuating asymmetry (FA) of morphological traits has attracted great attention as a short-cut measure of individual quality. Whereas there is some evidence that FA of sexual ornaments is negatively associated with immune function, studies concerning FA and immune function in non-ornamental traits are absent. Here, we tested whether FA of three non-ornamental traits in hind limbs is related to male immune function in a population of the Mediterranean field cricket, Gryllus bimaculatus. As different measures of male immune function, we used encapsulation rate and lytic activity. We found that a composite measure of FA (cFA) was negatively related to encapsulation rate. However, lytic activity was not related to cFA, but there was a tendency that males with higher body mass had higher lytic activity than males with lower body mass. Our results suggest that FA in non-ornamental traits indicates male immunocompetence in G. bimaculatus .     

Fluctuating asymmetry of leaves is a poor indicator of environmental stress and genetic stress by inbreeding in Silene vulgaris

Fluctuating asymmetry (FA) is often used as a measure of developmental instability and has been proposed as an indicator of both environmental and genetic stress. However, the empirical evidence for the effects of stress on FA in plants is inconsistent, and there are few controlled experimental studies. We analyzed different distance- and shape-based measures of the fluctuating asymmetry of the leaves of clonally replicated self- and cross-pollinated lineages of Silene vulgaris (Moench) Garcke (Caryophyllaceae) grown under a control and seven different stress treatments (drought, copper, simulated herbivory, and two levels of nutrient deficiency and of shade). Overall, FA differed among stress treatments, but was not generally higher under stress and even reduced in some treatments. Different measures of FA were only weakly correlated. Inbreeding increased only one measure of leaf FA, and only under high stress intensities. Our findings suggest that in S. vulgaris leaf fluctuating asymmetry does not serve as an indicator of environmental stress or of genetic stress by inbreeding.     

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.     

Fluctuating asymmetry and psychometric intelligence.

Little is known about the genetic nature of human psychometric intelligence (IQ), but it is widely assumed that IQ''s heritability is at loci for intelligence per se. We present evidence consistent with a hypothesis that interindividual IQ differences are partly due to heritable vulnerabilities to environmental sources of developmental stress, an indirect genetic mechanism for the heritability of IQ. Using fluctuating asymmetry (FA) of the body (the asymmetry resulting from errors in the development of normally symmetrical bilateral traits under stressful conditions), we estimated the relative developmental instability of 112 undergraduates and administered to them Cattell''s culture fair intelligence test (CFIT). A subsequent replication on 128 students was performed. In both samples, FA correlated negatively and significantly with CFIT scores. We propose two non-mutually exclusive physiological explanations for this correlation. First, external body FA may correlate negatively with the developmental integrity of the brain. Second, individual energy budget allocations and/or low metabolic efficiency in high-FA individuals may lower IQ scores. We review the data on IQ in light of our findings and conclude that improving developmental quality may increase average IQ in future generations.     

Fluctuating asymmetry does not reflect environmental stress during incubation in an oviparous lizard

Abstract

We analysed the relationship between incubation regime and fluctuating asymmetry in a cohort of juvenile lizards (Oligosoma suteri) incubated under laboratory conditions. We found no relationship between incubation temperature or water potential and the level of asymmetry in two cranial traits, although previous research showed profound effects of incubation temperature on gross morphology and hatching success. Our results suggest that fluctuating asymmetry is not a reliable indicator of developmental stress in this species.     

Fluctuating asymmetry as an ecological indicator of heavy metal stress in Lythrum salicaria

Fluctuating asymmetry (FA), a measure of random deviation of organismal traits from perfect bilateral symmetry, can be induced by environmental and/or genetic stress. We have conducted a replicated experiment to test the effect of a locally abundant heavy metal pollutant, lead, on the FA of a common wetland invasive species, Lythrum salicaria (purple loosestrife). The exposure to lead significantly reduced the shoot length, number and length of leaves as well as the biomass, but increased FA of L. salicaria leaves. The investigation suggests that FA in L. salicaria may be used as an ecological indicator to identify environmental stress caused by certain heavy metal pollutants. However, more studies are needed for the use of FA in L. salicaria as a comprehensive tool for biomonitoring of a wider array of environmental pollutants.     

Fluctuating dental asymmetry: Variation among skeletal populations

Recent investigations have shown that nongenetic, environmental factors can adversely affect dental growth and produce bilateral asymmetries in tooth size. When asymmetries do not favor either side, i.e., absence of directional asymmetry, the condition is termed fluctuating asymmetry. Fluctuating asymmetry of the mesiodistal and buccolingual dimensions of the total permanent dentition was compared among human skeletal populations which differ socio-economically and nutritionally. Odontometric data were collected from prehistoric hunters (Indian Knoll site), later aboriginal farming groups (Campbell and Larson sites), and a modern cadaver population (Hamann-Todd). The magnitude of asymmetry is expressed by the familiar correlation coefficient, r. The proportion, then, of intra-individual variation due to fluctuating asymmetry is equal to 1-r. With Wilcoxon's signed ranks test on the correlation coefficients no significant sex difference was shown within populations. Among groups, though, Indian Knoll was the most odontometrically asymmetrical; moreover, within Indian Knoll, the taller and ostensibly better nourished individuals had larger, less asymmetrical teeth than the shorter individuals. These results suggest that environmentally mediated growth disturbance may be sensitively reflected by dental asymmetry. A population exhibiting other signs of severe growth disturbance, e.g., enamel hypoplasia and Harris lines, was the most dentally asymmetrical.     

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.     

Fluctuating asymmetry in a secondary sexual trait: no associations with individual fitness, environmental stress or inbreeding, and no heritability

It has been suggested that fluctuating asymmetry (FA) in secondary sexual traits may be a useful indicator of either individual quality or environmental stress. We tested this concept using a series of analyses of FA in male antler size in a wild red deer (Cervus elaphus) population, using four measures of size repeated across successive years on the same individuals. We found no consistent evidence of correlations between traits in levels of FA, nor of any associations between known environmental or developmental conditions. None of the four measures of FA showed a significant heritability (average h2 = 0.041), nor was there any evidence of inbreeding depression. For three of the four traits, fluctuating asymmetry did not predict either annual or lifetime breeding success. However there were significant associations between breeding success and FA in antler length. Given the series of null results in our other tests, it seems likely that this was a direct mechanistic effect rather than because measures of FA were indicative of individual quality or condition.