The ecological stress theory of aging and hormesis: an energetic evolutionary model.

Free-living organisms normally struggle to exist in harsh environments that are nutritionally and energetically inadequate, where evolutionary adaptation is challenged by internal stresses within organisms and external stresses from the environment. The incorporation of environmental variables into aging theories such as the free-radical and metabolic rate/oxidative stress theories, is the basis of the ecological stress theory of aging and hormesis. Environmental variation from optimum to lethal extremes gives a fitness-stress continuum, where energetic efficiency, or fitness, is inversely related to stress level; in the evolutionary context survival is a more direct measure of fitness for assessing aging than is lifespan. On this continuum, the hormetic zone is in the optimum region, while aging emphasizes survival towards lethal extremes. At the limits of survival, a convergence of physiological and genetical processes is expected under accumulating stress from Reactive Oxygen Species, ROS. Limited ecologically-oriented studies imply that major genes are important towards limits of survival compared with the hormetic zone. Future investigations could usefully explore outlier populations physiologically and genetically, since there is the likelihood that genetic variability may be lower in those cohorts managing to survive to extremely advanced ages as found in highly stressed ecological outlier populations. If so, an evolutionary explanation of the mortality-rate decline typical of cohorts of the extremely old emerges. In summary, an energetic evolutionary approach produces a general aging theory which automatically incorporates hormesis, since the theory is based on a fitness-stress continuum covering the whole range of possible abiotic environments of natural populations.