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.

Fluctuating asymmetry, the random deviation from perfect symmetry, is a widely used population-level index of developmental instability, developmental noise, and robustness. It reflects a population’s state of adaptation and genomic coadaptation. Here, we review the literature on fluctuating asymmetry of human populations. The most widely used bilateral traits include skeletal, dental, and facial dimensions; dermatoglyphic patterns and ridge counts; and facial shape. Each trait has its advantages and disadvantages, but results are most robust when multiple traits are combined into a composite index of fluctuating asymmetry (CFA). Both environmental (diet, climate, toxins) and genetic (aneuploidy, heterozygosity, inbreeding) stressors have been linked to population-level variation in fluctuating asymmetry. In general, these stressors increase average fluctuating asymmetry. Nevertheless, there have been many conflicting results, in part because (1) fluctuating asymmetry is a weak signal in a sea of noise; and (2) studies of human fluctuating asymmetry have not always followed best practices. The most serious concerns are insensitive asymmetry indices (correlation coefficient and coefficient of indetermination), inappropriate size scaling, unrecognized mixture distributions, inappropriate corrections for directional asymmetry, failure to use composite indices, and inattention to measurement error. Consequently, it is often difficult (or impossible) to compare results across traits, and across studies.

Multiplicative error accounts for much of the size-scaling and leptokurtosis in fluctuating asymmetry. It arises when growth involves the addition of