The Evolution of Dispersal in Random Environments and The Principle of Partial Control

McNamara and Dall (2011) identified novel relationships between the abundance of a species in different environments, the temporal properties of environmental change, and selection for or against dispersal. Here, the mathematics underlying these relationships in their two-environment model are investigated for arbitrary numbers of environments. The effect they described is quantified as the fitness-abundance covariance. The phase in the life cycle where the population is censused is crucial for the implications of the fitness-abundance covariance. These relationships are shown to connect to the population genetics literature on the Reduction Principle for the evolution of genetic systems and migration. Conditions that produce selection for increased unconditional dispersal are found to be new instances of departures from reduction described by the "Principle of Partial Control" proposed for the evolution of modifier genes. According to this principle, variation that only partially controls the processes that transform the transmitted information of organisms may be selected to increase these processes. Mathematical methods of Karlin, Friedland, and Elsner, Johnson, and Neumann, are central in generalizing the analysis. Analysis of the adaptive landscape of the model shows that the evolution of conditional dispersal is very sensitive to the spectrum of genetic variation the population is capable of producing, and suggests that empirical study of particular species will require an evaluation of its variational properties.Comment: Dedicated to the memory of Professor Michael Neumann, one of whose many elegant theorems provides for a result presented here. 28 pages, 1 table, 1 figur

Fundamental Properties of the Evolution of Mutational Robustness

Evolution on neutral networks of genotypes has been found in models to concentrate on genotypes with high mutational robustness, to a degree determined by the topology of the network. Here analysis is generalized beyond neutral networks to arbitrary selection and parent-offspring transmission. In this larger realm, geometric features determine mutational robustness: the alignment of fitness with the orthogonalized eigenvectors of the mutation matrix weighted by their eigenvalues. "House of cards" mutation is found to preclude the evolution of mutational robustness. Genetic load is shown to increase with increasing mutation in arbitrary single and multiple locus fitness landscapes. The rate of decrease in population fitness can never grow as mutation rates get higher, showing that "error catastrophes" for genotype frequencies never cause precipitous losses of population fitness. The "inclusive inheritance" approach taken here naturally extends these results to a new concept of dispersal robustness.Comment: 17 pages, 1 figur

Simon-Ando decomposability and fitness landscapes

In this paper, we investigate fitness landscapes (under point mutation and recombination) from the standpoint of whether the induced evolutionary dynamics have a “fast-slow” time scale associated with the differences in relaxation time between local quasi-equilibria and the global equilibrium. This dynamical hevavior has been formally described in the econometrics literature in terms of the spectral properties of the appropriate operator matrices by Simon and Ando (Econometrica 29 (1961) 111), and we use the relations they derive to ask which fitness functions and mutation/recombination operators satisfy these properties. It turns out that quite a wide range of landscapes satisfy the condition (at least trivially) under point mutation given a sufficiently low mutation rate, while the property appears to be difficult to satisfy under genetic recombination. In spite of the fact that Simon-Ando decomposability can be realized over fairly wide range of parameters, it imposes a number of restriction on which landscape partitionings are possible. For these reasons, the Simon-Ando formalism does not appear to be applicable to other forms of decomposition and aggregation of variables that are important in evolutionary systems

Proof of the Feldman-Karlin Conjecture on the Maximum Number of Equilibria in an Evolutionary System

Feldman and Karlin conjectured that the number of isolated fixed points for deterministic models of viability selection and recombination among n possible haplotypes has an upper bound of 2^n - 1. Here a proof is provided. The upper bound of 3^{n-1} obtained by Lyubich et al. (2001) using Bezout's Theorem (1779) is reduced here to 2^n through a change of representation that reduces the third-order polynomials to second order. A further reduction to 2^n - 1 is obtained using the homogeneous representation of the system, which yields always one solution `at infinity'. While the original conjecture was made for systems of viability selection and recombination, the results here generalize to viability selection with any arbitrary system of bi-parental transmission, which includes recombination and mutation as special cases. An example is constructed of a mutation-selection system that has 2^n - 1 fixed points given any n, which shows that 2^n - 1 is the sharpest possible upper bound that can be found for the general space of selection and transmission coefficients.Comment: 9 pages, 1 figure; v.4: final minor revisions, corrections, additions; v.3: expands theorem to cover all cases, obviating v.2 distinction of reducible/irreducible; details added to: discussion of Lyubich (1992), example that attains upper bound, and homotopy continuation method