FLUCTUATING ASYMMETRY: Measurement, Analysis, Patterns

With these words Darwin opened a brief paragraph citing observations antithetical to his supposition: anecdotal reports of the inheritance of characters missing from one side of the body. His initial hunch, however, has stood the test of time: Genetic studies have confirmed that where only small, random deviations from bilateral symmetry exist, the deviations in a particular direction have little or no measurable heritability (17, 47, 51, 65a, 73a, 74, 91). The genetic basis of bilateral symmetry thus appears to differ fundamentally from that of virtually all other morphological features. Why, then, all the recent (and not so recent) interest in such minor, nondirectional deviations from bilateral symmetry [fluctuating asymmetry (FA); 60 cited in 99]? Four reasons. First, FA relates in a curious way to what is perhaps the major unsolved general problem in modem biology: the orderly expression of genotypes as complex, three-dimensional phenotypes. As was emphasized in a flurry of activity in the mid to late 1950s, and many times since, FA provides an appealing measure of 'developmental noise,' or minor environmentally induced departures from some ideal developmental program (101). Its appeal derives from an a priori knowledge of the ideal: perfect bilateral symmetry. For unilateral characters, the ideal is rarely known (but see 1, 2, and 59 for one possible approach). A second reason for interest in