Developmental integration and evolution of labile plasticity in a complex quantitative character in a multiperiodic environment.

Labile plasticity in a complex quantitative character is modeled, with multiple components contributing to net plasticity in the character. Each component has a specific development rate, norm of reaction, and cost of plasticity. For example, thermal adaptation in mammals includes seasonal fat deposition and fur growth, short-term shivering and sweating or panting, and movement between warm and cold sites. Norms of reaction do not reveal patterns of developmental integration, which must be investigated by studies of developmental dynamics in a changing environment. In a periodic environment, a labile character with a single component of plasticity is constrained by filtering environmental frequencies above the development rate and by the cost of plasticity. With multiple components of plasticity, some patterns of integration can alleviate these constraints to greatly improve fidelity of the mean phenotype tracking multiperiodic cycles in the optimum phenotype. This occurs by environmental signal amplification or inhibition through developmental integration among components and by an augmented development rate of net plasticity in the character that reduces environmental frequency filtering. When development of a component with high cost of plasticity is regulated partly by the norm of reaction of another component, evolution can diminish the reaction norm slope of the costly component without curtailing its development, thereby reducing the loss of fitness from its cost of plasticity. Apparent maladaptation in a component of plasticity may be an integral part of an adaptive pattern of developmental integration by mutual inhibition between components and compensatory evolution of a negative component reaction norm slope.