Nutritional ecology and the evolution of aging.

Considerable progress has been made in understanding both evolutionary and mechanistic aspects of biological aging, although the two areas remain poorly integrated. We suggest that a greater emphasis on ecology can help to remedy this, by focusing on the interface between biological mechanisms and the environments in which they evolved by natural selection. Among the most salient aspects of the environment relevant to aging is nutrition, and yet in the bulk of aging research nutrition is coarsely represented as dietary restriction or caloric restriction, without consideration for how specific components of diet, beyond "energy" (the undifferentiated mix of macronutrients), are driving the observed effects. More recently, it has become clear that specific nutrients (notably amino acids) and interactions among nutrients (i.e., nutritional balance) play important roles in the biology of aging. We show how a method developed in nutritional ecology, called the Geometric Framework for nutrition, can help to understand the nutritional interactions of animals with their environments, by explicitly distinguishing the roles of calories, individual nutrients and nutrient balance. Central to these models are the active regulatory responses that animals use to mediate between variation in the nutritional environment and fitness-related consequences such as lifespan and reproduction. These homeostatic responses provide a guide for researchers that can help to link the biological mechanisms with evolutionary processes in the context of a multi-dimensional nutritional environment.