Life not lived due to disequilibrium in heterogeneous age-structured populations.

Three models of age-structured populations with demographically heterogeneous subpopulations are analyzed. In the first model, each subpopulation has its own age-specific vital rates which are fixed in time. In the second model, the vital rates of each subpopulation are uniformly inhibited by increasing total numbers of individuals. In the third, the vital rates of groups of subpopulations are inhibited by the total numbers of individuals in other groups of subpopulations with an intensity that depends on the interacting pair of groups. Three functions are defined to measure disequilibrium in the subpopulation frequencies, subpopulation age structures, and total population size. For the first model, we show that disequilibrium will shift the trajectory of the total numbers of individuals forward or backward in time by an asymptotic constant that is proportional to the sum of the disequilibrium measures. For the second model, we establish sufficient conditions for the existence of a globally stable equilibrium and we show that disequilibrium will result in a finite loss or gain in life which is proportional to the sum of the disequilibrium measures. For the last model, we show that the loss or gain in life for each group of subpopulations is a linear combination over all groups of the sums of the three disequilibrium measures. We illustrate these results with numerical examples and give possible biological interpretations of the models. We relate these new results to previous work on the cost of natural selection and measures of demographic disequilibrium.