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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.

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.

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.

Adaptive developmental plasticity allows individuals to match their phenotype with their environment, increasing fitness where threats are inconsistently present. However, despite clear advantages of plasticity, adaptive traits are not ubiquitously nor infinitely plastic. Trade-offs between benefits and costs or limits are therefore theoretically necessary to constrain the evolution of plastic responses. Systems in which extreme risk can be reliably detected are ideal for investigating trade-offs, as even costly responses may be adaptive where risk is severe. Cane toads (Rhinella marina) are abundant in Australia and produce large clutches (frequently >10,000 eggs), but asynchronous breeding and rapid development result in variable larval densities within breeding pools. In the field, we found that cannibalism by older cohorts often reduces the survival of conspecific eggs and newly hatched pre-feeding larvae (\"hatchlings\") by >99%, as feeding larvae (\"tadpoles\") use chemical cues from the relatively immobile hatchlings to locate and consume them. After hatchlings become free-swimming, however, they cannot be cannibalized. Hatchlings can reduce this period of vulnerability by accelerating development when they detect cannibal cues. However, this developmental acceleration decreases initial tadpole mass, reduces subsequent survival, growth, and development, affects behavior, and compromises feeding structures. Reaction norms differ among clutches, and greater developmental acceleration is followed by greater impairment of larval function in plastic clutches, whereas nonresponsive clutches are unaffected by cue exposure. More plastic clutches ultimately exhibit both poorer performance and greater variation among siblings in exposed and (to a lesser degree) control treatments. Variation among clutches in tadpole viability is driven by differences in plasticity rather than phenotype; fitness reductions are linked to developmental acceleration, not rapid development per se. Clutches with intrinsically slow prefeeding developmental rates exhibit stronger acceleration (i.e., steeper reaction norms), but clutches with intrinsically rapid development reach invulnerable stages more quickly than those that accelerate development. As a result, high cannibalism risk may favor canalized rapid development rather than facultative developmental acceleration. Cannibalism plays an important role in the recruitment of this invasive species, and hatchling defenses against this threat demonstrate how the limits and costs associated with an inducible defense can favor canalized defenses over phenotypic plasticity.

In many species, including humans and Drosophila, symmetric individuals secure more matings, suggesting that bilateral symmetry signals the quality of potential mates and is subject to sexual selection. However, this idea remains controversial, largely because obtaining conclusive experimental evidence has been hindered by confounding effects arising from the methods used to increase asymmetry in test subjects. Here, we show that altering gravity during development increases asymmetry in Drosophila melanogaster without a detrimental effect on survival, growth, and behavior. Testing males with altered-gravity–induced asymmetry in female mate-choice assays revealed symmetry-based discrimination of males via auditory cues. Females similarly discriminated against males with genetically induced asymmetry, suggesting that their preference for symmetry is not specific to altered gravity. By segmenting the male courtship song into left and right wing-generated song-bouts, we detected asymmetry in the courtship song of altered-gravity males with asymmetric wings that experienced rejection. Females experimentally evolved in the absence of mate choice lacked this preference for symmetry, suggesting that symmetry is maintained by sexual selection. Our data provide evidence for the role of symmetry in sexual selection and reveal how nonvisual cues can flag mate asymmetry during courtship.

Phenotypic variation arises from genetic and environmental variation, as well as random aspects of development. The genetic (nature) and environmental (nurture) components of this variation have been appreciated since at least 1900. The random developmental component (noise) has taken longer for quantitative geneticists to appreciate. Here, I sketch the historical development of the concepts of random developmental noise and developmental instability, and its quantification via fluctuating asymmetry. The unsung pioneers in this story are Hugo DeVries (fluctuating variation, 1909), C. H. Danforth (random variation between monozygotic twins, 1919), and Sewall Wright (random developmental variation in piebald guinea pigs, 1920). The first pioneering study of fluctuating asymmetry, by Sumner and Huestis in 1921, is seldom mentioned, possibly because it failed to connect the observed random asymmetry with random developmental variation. This early work was then synthesized by Boris Astaurov in 1930 and Wilhelm Ludwig in 1932, and then popularized by Drosophila geneticists beginning with Kenneth Mather in 1953. Population phenogeneticists are still trying to understand the origins and behavior of random developmental variation. Some of the developmental noise represents true stochastic behavior of molecules and cells, while some represents deterministic chaos, nonlinear feedback, and symmetry breaking.

As natural disasters become more frequent due to climate change, understanding the biological impact of these ecological catastrophes on wild populations becomes increasingly pertinent. Fluctuating asymmetry (FA), or random deviations from bilateral symmetry, is reflective of developmental instability and has long been positively associated with increases in environmental stress. This study investigates craniofacial FA in a population of free‐ranging rhesus macaques (Macaca mulatta) that has experienced multiple Category 3 hurricanes since the colony's inception on Cayo Santiago, including 275 individuals from ages 9 months to 31 years (F = 154; M = 121). Using geometric morphometrics to quantify FA and a linear mixed‐effect model for analysis, we found that sex, age, and decade of birth did not influence the amount of FA in the individuals included in the study, but the developmental stage at which individuals experienced these catastrophic events greatly impacted the amount of FA exhibited (p = .001). Individuals that experienced these hurricanes during fetal life exhibited greater FA than any other post‐natal developmental period. These results indicate that natural disasters can be associated with developmental disruption that results in long‐term effects if occurring during the prenatal period, possibly due to increases in maternal stress‐related hormones. Resumen A medida que los desastres naturales se vuelven más frecuentes debido al cambio climático, entender el impacto biológico de estas catástrofes ecológicas en poblaciones silvestres va en aumento pertinente. La asimetría fluctuante (AF), o desviaciones aleatorias de simetría bilateral, es reflejo de inestabilidad durante el desarrollo y se ha asociado positivamente con incrementos en estrés ambiental durante mucho tiempo. Este estudio investiga AF craneofacial en una población de macacos rhesus (Macaca mulatta) en libertad que ha experimentado múltiples huracanes categoría 3 desde el inicio de la colonia en Cayo Santiago, e incluye 275 individuos de 9 meses a 31 años de edad (F = 154; M = 121). Usando morfometría geométrica para cuantificar AF y un modelo lineal de efectos mixtos para análisis, encontramos que el sexo, la edad y la década de nacimiento no influyeron en la cantidad de AF en los individuos incluidos en el estudio, pero la etapa de desarrollo en la que los individuos experimentaron estos eventos catastróficos impactó altamente la cantidad de AF exhibida (p = .001). Los individuos que experimentaron estos huracanes durante el período fetal exhibieron mayor AF que cualquier otro período de desarrollo posnatal. Estos resultados indican que los desastres naturales pueden asociarse con trastornos del desarrollo que tienen efectos a largo plazo si ocurren durante el período prenatal, posiblemente debido al aumento de hormonas maternas relacionadas con el estrés. Fluctuating asymmetry (FA), or random deviations from symmetry, can be used as a proxy for developmental instability, and anthropogenic and natural disruptions have been associated with increased levels of FA. In this study, we investigate the effect of demographic variables (age, sex, and decade of birth) and hurricane experience on FA in a model primate taxon: rhesus macaques (Macaca mulatta) to better understand the impact major ecological catastrophes have on skeletal development. Individuals that experienced a hurricane event in utero exhibited higher levels of FA than juvenile or adult individuals that experienced that same event, which could be due to increased maternal stress hormones such as glucocorticoids that can disrupt fetal development.

Developmental instability (DI) is thought to be inversely related to a capacity of an organism to buffer its development against random genetic and environmental perturbations. DI is represented by a trait’s inter- and intra-individual variabilities. The inter-individual variability (inversely referred to as canalization) indicates the capability of organisms to reproduce a trait from individual to individual. The intra-individual variability reflects an organism’s capability to stabilize a trait internally under the same conditions, and, for symmetric traits, it is expressed as fluctuating asymmetry (FA). When representing a trait as a random variable conditioned on environmental fluctuations, it is clear that, in statistical terms, the DI partitions into “extrinsic” (canalization) and “intrinsic” (FA) components of a trait’s variance/noise. We established a simple statistical framework to dissect both parts of a symmetric trait variance/noise using a PCA (principal component analysis) projection of the left/right measurements on eigenvectors followed by GAMLSS (generalized additive models for location scale and shape) modeling of eigenvalues. The first eigenvalue represents “extrinsic” and the second—“intrinsic” DI components. We applied this framework to investigate the impact of mother–fetus major histocompatibility complex (MHC)-mediated immune cross-talk on gene expression noise and developmental stability. We showed that “intrinsic” gene noise for the entire transcriptional landscape could be estimated from a small subset of randomly selected genes. Using a diagnostic set of genes, we found that allogeneic MHC combinations tended to decrease “extrinsic” and “intrinsic” gene noise in C57BL/6J embryos developing in the surrogate NOD-SCID and BALB/c mothers. The “intrinsic” gene noise was negatively correlated with growth (embryonic mass) and the levels of placental growth factor (PLGF), but not vascular endothelial growth factor (VEGF). However, it was positively associated with phenotypic growth instability and noise in PLGF. In mammals, the mother–fetus MHC interaction plays a significant role in development, contributing to the fitness of the offspring. Our results demonstrate that a positive impact of distant MHC combinations on embryonic growth could be mediated by the reduction of “intrinsic” gene noise followed by the developmental stabilization of growth.

The eco-evolutionary theory of developmental instability predicts that small, non-directional deviations from perfect symmetry in morphological traits (termed fluctuating asymmetry, FA) emerge when an individual is unable to buffer environmental or genetic stress during its development. Consequently, FA is widely used as an index of stress. The goal of the present study was to experimentally test a seemingly trivial prediction derived from the theory of developmental instability—and from previous observational studies—that significant growth retardation (which indicates stress) in plants is accompanied by an increase in FA of their leaves. We induced stress, evidenced by a significant decrease in biomass relative to control, in cucumber (Cucumis sativus), sweet pepper (Capsicum annuum), and common bean (Phaseolus vulgaris) by applying water solutions of copper and nickel to the soil in which plants were grown. Repeated blind measurements of plant leaves revealed that leaf FA did not differ between stressed and control plants. This finding, once again, demonstrated that FA cannot be seen as a universal indicator of environmental stress. We recommend that the use of FA as a stress index is discontinued until the scope of the developmental instability theory is clarified and its applicability limits are identified.

A key focus of evolutionary developmental biology is on how phenotypic diversity is generated. In particular, both plasticity and developmental instability contribute to phenotypic variation among genetically identical individuals, but the interactions between the two phenomena and their general fitness impacts are unclear. We discovered a striking example of asymmetry in pea aphids: the presence of wings on one side and the complete or partial absence of wings on the opposite side. We used this asymmetric phenotype to study the connection between plasticity, developmental instability and fitness. We found that this asymmetric wing development (i) occurred equally on both sides and thus is a developmental instability; (ii) is present in some genetically unique lines but not others, and thus has a genetic basis; and (iii) has intermediate levels of fecundity, and thus does not necessarily have negative fitness consequences. We conclude that this dramatic asymmetry may arise from incomplete switching between developmental targets, linking plasticity and developmental instability. We suspect that what we have observed may be a more widespread phenomenon, occurring across species that routinely produce distinct, alternative phenotypes.