Approximately two decades after the first pioneering analyses, the study of shape asymmetry with the methods of geometric morphometrics has matured and is a burgeoning field. New technology for data ...collection and new methods and software for analysis are widely available and have led to numerous applications in plants and animals, including humans. This review summarizes the concepts and morphometric methods for studying asymmetry of shape and size. After a summary of mathematical and biological concepts of symmetry and asymmetry, a section follows that explains the methods of geometric morphometrics and how they can be used to analyze asymmetry of biological structures. Geometric morphometric analyses not only tell how much asymmetry there is, but also provide information about the patterns of covariation in the structure under study. Such patterns of covariation in fluctuating asymmetry can provide valuable insight about the developmental basis of morphological integration, and have become important tools for evolutionary developmental biology. The genetic basis of fluctuating asymmetry has been studied from empirical and theoretical viewpoints, but serious challenges remain in this area. There are many promising areas for further research that are only little explored at present.
Fluctuating asymmetry (FA), in contrast with other asymmetries, is the bilateral asymmetry that represents small, random developmental differences between right and left sides. After nearly a century ...of using traditional morphometrics in the estimation of FA, geometric morphometrics (GM) now provides new insights into the use of FA as a tool, especially for assessing environmental and developmental stress. Thus, it will be possible to assess adaptation to various environmental stressors as particular triggers for unavoidable selection pressures. In this review, we describe measures of FA that use geometric morphometrics, and we include a flow chart of the methodology. We also describe how this combination (GM + FA) has been tested in several agroecosystems. Nutritional stress, temperature, chemical pollution, and population density are known stressors experienced by populations in agroecosystems.
•Hallucinations and disorganisation were associated with increased body asymmetry.•The schizophrenia patients without these symptoms presented normal body asymmetry.•In healthy individuals body ...asymmetry was associated with propensity to anxiety.
Fluctuating asymmetry represents the degree to which the right and left side of the body are asymmetrical, and is a sign of developmental instability. Higher levels of fluctuating asymmetry have been observed in individuals within the schizophrenia spectrum. We aimed to explore the associations of fluctuating asymmetry with psychotic and affective symptoms in schizophrenia patients, as well as with propensity to these symptoms in non-clinical individuals. A measure of morphological fluctuating asymmetry was calculated for 39 patients with schizophrenia and 60 healthy individuals, and a range of clinical and subclinical psychiatric symptoms was assessed. Regression analyses of the fluctuating asymmetry measure were conducted within each group. In the patient cohort, fluctuating asymmetry was significantly associated with the hallucination and thought disorganisation scores. T-test comparisons revealed that the patients presenting either hallucinations or thought disorganisation were significantly more asymmetrical than were the healthy individuals, while the patients without these key symptoms were equivalent to the healthy individuals. A positive association with the anxiety score emerged in a subsample of 36 healthy participants who were rated on affective symptoms. These findings suggest that fluctuating asymmetry may be an indicator of clinical hallucinations and thought disorganisation rather than an indicator of schizophrenia disease.
Adaptive responses to autocorrelated environmental fluctuations through evolution in mean reaction norm elevation and slope and an independent component of the phenotypic variance are analyzed using ...a quantitative genetic model. Analytic approximations expressing the mutual dependencies between all three response modes are derived and solved for the joint evolutionary outcome. Both genetic evolution in reaction norm elevation and plasticity are favored by slow temporal fluctuations, with plasticity, in the absence of microenvironmental variability, being the dominant evolutionary outcome for reasonable parameter values. For fast fluctuations, tracking of the optimal phenotype through genetic evolution and plasticity is limited. If residual fluctuations in the optimal phenotype are large and stabilizing selection is strong, selection then acts to increase the phenotypic variance (bethedging adaptive). Otherwise, canalizing selection occurs. If the phenotypic variance increases with plasticity through the effect of microenvironmental variability, this shifts the joint evolutionary balance away from plasticity in favor of genetic evolution. If microenvironmental deviations experienced by each individual at the time of development and selection are correlated, however, more plasticity evolves. The adaptive significance of evolutionary fluctuations in plasticity and the phenotypic variance, transient evolution, and the validity of the analytic approximations are investigated using simulations.
Stress experienced during development in organisms with bilateral structures could result in developmental instability, which is expressed as subtle non‐directional deviations from perfect symmetry, ...known as fluctuating asymmetry (FA). As such, FA has been proposed, and extensively used, as a trait indicating stress for many organisms with bilateral structures and many types of stress. However, while this concept may apply to animals, the evidence for plants' main vegetative structures, that is, leaves, remains equivocal, and a comprehensive synthesis on this topic is still missing.
We designed observational field and controlled greenhouse studies, combining different growth and leaf forms across multiple stress gradients, comprising 21 species and 80 populations. We measured FA as the difference between the left and right area of the leaf, an approach that accommodates diverse leaf forms. We used high‐precision, blind, single‐person measurements and tested for other forms of symmetry and the effect of leaf size. We further complemented our study with a systematic literature review of FA in plant leaves, compiling 51 studies comprising 72 species, 23 stress types and 131 unique entries (species × stress type).
We consistently found no effect of stress on leaf FA in any of the studied species in both our field and experimental gradients. In the systematic literature review, only 39% of the unique entries showed the expected increase in FA with stress, 53% showed no effect, 9% showed an opposite trend of a decrease in FA with stress and 1% showed a unimodal relationship. Importantly, only 40% of all entries fulfilled the crucial step of controlling for a high‐precision measurement, and of these 49% reported the expected increase in FA with stress.
Both the results of our observational and experimental approaches and the systematic literature review failed to support a clear relationship between stress and FA in plant leaves. These results clearly show that FA in plant leaves cannot be used as a reliable trait indicating stress during development.
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Read the free Plain Language Summary for this article on the Journal blog.
•Fluctuating asymmetry (FA) is a less invasive indicator of effect of stress.•Treatments had an effect on femoral pore but non on head shape FA.•A dissimilar degree of FA among treatments was found ...only in females.•FA may have a real potential as biomarker of population stress in wall lizards.
The extensive use of pesticides in agricultural environments produces drastic effects on wildlife, hence the need for less invasive indicators of environmental stress to monitoring the impact of agriculture treatments on biological systems. Fluctuating asymmetry (FA), as measure of developmental instability, has recently been proposed as reliable biomarker of populations stress due to environmental disturbance. We investigate femoral pores (FP) and dorsal head shape (HS) traits in populations of the Italian wall lizard inhabiting agricultural environments to examine whether different pesticide exposures (conventional, organic and control) can cause distinctive degree of FA. High-resolution photographs of FP and HS were taken in the field with a digital camera. The number of FP were counted twice on both sides and HS was analysed using geometric morphometrics with 25 landmarks and 12 semilandmarks. Individuals under conventional management showed higher levels of FA compared to control ones, and females exhibited higher FA levels than males for the FP. However, no significant difference was found for the HS trait. Our study provided evidence that FA may have a real potential as biomarker of population stress in wall lizards, highlighting the importance in the choice of the experimental design and the traits adopted for estimating DI.
Best practices in studies of developmental instability, as measured by fluctuating asymmetry, have developed over the past 60 years. Unfortunately, they are haphazardly applied in many of the papers ...submitted for review. Most often, research designs suffer from lack of randomization, inadequate replication, poor attention to size scaling, lack of attention to measurement error, and unrecognized mixtures of additive and multiplicative errors. Here, I summarize a set of best practices, especially in studies that examine the effects of environmental stress on fluctuating asymmetry.
Fluctuating asymmetry consists of random deviations from perfect symmetry in populations of organisms. It is a measure of developmental noise, which reflects a population’s average state of ...adaptation and coadaptation. Moreover, it increases under both environmental and genetic stress, though responses are often inconsistent. Researchers base studies of fluctuating asymmetry upon deviations from bilateral, radial, rotational, dihedral, translational, helical, and fractal symmetries. Here, we review old and new methods of measuring fluctuating asymmetry, including measures of dispersion, landmark methods for shape asymmetry, and continuous symmetry measures. We also review the theory, developmental origins, and applications of fluctuating asymmetry, and attempt to explain conflicting results. In the process, we present examples from the literature, and from our own research at “Evolution Canyon” and elsewhere.
Developmental instability in domesticated mammals Wilson, Laura A. B.
Journal of experimental zoology. Part B, Molecular and developmental evolution,
December 2022, Letnik:
338, Številka:
8
Journal Article
Recenzirano
Measures of fluctuating asymmetry (FA) have been adopted widely as an estimate of developmental instability. Arising from various sources of stress, developmental instability is associated with an ...organism's capacity to maintain fitness. The process of domestication has been framed as an environmental stress with human‐specified parameters, suggesting that FA may manifest to a larger degree among domesticates compared to their wild relatives. This study used three‐dimensional geometric morphometric landmark data to (a) quantify the amount of FA in the cranium of six domestic mammal species and their wild relatives and, (b) provide novel assessment of the commonalities and differences across domestic/wild pairs concerning the extent to which random variation arising from the developmental system assimilates into within‐population variation. The majority of domestic mammals showed greater disparity for asymmetric shape, however, only two forms (Pig, Dog) showed significantly higher disparity as well as a higher degree of asymmetry compared to their wild counterparts (Wild Boar, Wolf). Contra to predictions, most domestic and wild forms did not show a statistically significant correspondence between symmetric shape variation and FA, however, a moderate correlation value was recorded for most pairs (r‐partial least squares >0.5). Within pairs, domestic and wild forms showed similar correlation magnitudes for the relationship between the asymmetric and symmetric components. In domesticates, new variation may therefore retain a general, conserved pattern in the gross structuring of the cranium, whilst also being a source for response to selection on specific features.
RESEARCH HIGHLIGHTS
The majority of domestic mammals showed greater disparity for asymmetric cranial shape, however, only two forms (Pig, Dog) showed significantly higher disparity as well as a higher degree of asymmetry compared to their wild counterparts (Wild Boar, Wolf).
Domestic mammals and their wild counterparts showed similar correlation magnitudes for the relationship between the asymmetric and symmetric components, suggesting that in domesticates new variation retains a general, conserved pattern in the gross structuring of the cranium, whilst also being a source for response to selection on specific features.
The process of domestication has been framed as an environmental stress with human‐specified parameters, suggesting that fluctuating asymmetry may manifest to a larger degree among domesticates compared to their wild relatives. In this study, I quantify the magnitude of cranial fluctuating asymmetry in pairs of wild and domestic mammals and assess how symmetric and asymmetric variation is patterned.
Random and subtle deviations from bilateral symmetry (fluctuating asymmetry) have long been of interest to biologists who wish to study the susceptibility of organisms to changes in environmental ...quality. However, the reliability of FA as a biomarker has come under question due to inconsistent results in the literature. We conducted a meta-analysis of published literature to test the hypothesis that FA is a reliable biomarker of environmental stress in insects and identify possible sources of variation amongst studies. We expected studies to detect larger, positive magnitudes of effect on FA in lab populations due to the lack of confounding effects from other environmental factors compared to wild populations. Additionally, we predicted that studies that used geometric morphometric approaches to FA in shape and size would be more sensitive to changes in environmental quality compared to linear and meristic measures and thus show larger effects on FA. We also expected anthropogenic stressors to generate significantly larger effects on FA compared to naturally occurring stressors due to the organisms’ inability to buffer developmental pathways against a novel stressor. Finally, we predicted comparatively larger magnitudes of effect in studies that verified the environmental factor acting on the organism was a stressor by detecting negative effects on fitness-related traits. Overall, we found that FA is a sensitive biomarker of environmental stress. Environmental stressors explained 36% of the variation of effect on FA across studies. Studies that demonstrated a negative effect of the stressor on fitness-related traits showed significantly larger, positive magnitudes of effect on FA compared to studies that did not detect an effect from the environmental stressor. Additionally, studies conducted under laboratory conditions detected significantly larger, effects on FA compared to field-based studies. The kind of trait measured and the novelty of the stressor did not significantly account for differences amongst studies. Thus, the use of FA as a biomarker of environmental stress is a legitimate tool particularly when studies verify the biological relevance of stressors for the study organism.
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