Patterns of growth and fluctuating asymmetry: the effects of asymmetrical investment in traits with determinate growth

Fluctuating asymmetry (FA, small and random deviations from perfect symmetry in otherwise bilaterally symmetrical traits) is assumed to depend directly on developmental stability (DS). However, since the processes related to DS operate at a microscopic level, the translation of microscopic errors to the macroscopic character asymmetries (i.e. the DS-FA relation) could be mediated by the cost of growth. To investigate the consequences of this hypothesis, I constructed a model with the following assumptions: the degree of development of a trait depends on the amount of resources allocated to growth, which may differ between individuals and between traits; the cost of growth increases with development, and developmental noise causes random asymmetries in biosynthesis in the two sides of the body. The model predicts that FA follows a parabolic trajectory as growth investment levels increase, regardless of DS. However, depending on the range of variation of investment levels, different relationships (positive, negative or flat) between trait size and asymmetry could be expected. This model may also explain why FA is correlated with the degree of stress in some traits but not in others, and why some traits present a higher level of FA than others. The model also suggests that FA measured on growing traits, at a certain size, could be a more reliable indicator of DS than FA measured on fully grown traits. A study analysing FA patterns of developing and fully grown feathers in the house sparrow (Passer domesticus) corroborates that the levels of FA, and the relationship between asymmetry and feather length, depend on the degree of feather development.