Adaptation and Natural Selection revisited

In Adaptation and Natural Selection, George C. Williams linked the distinction between group and individual adaptation with the distinction between group and individual selection. Williams’ Principle, as we will call it, says that adaptation at a level requires selection at that level. This is a necessary but not a sufficient condition; for example, group adaptation requires group selection, but the fact that group selection influences a trait’s evolution does not suffice for the resulting trait frequency to be a group adaptation. What more is required? In this paper, we describe an answer to this question that has been developed in multilevel selection theory. We also discuss an alternative framework for defining units of adaptation that violates Williams’ Principle.     

Natural Selection and Y-Linked Polymorphism

Several population genetic models allowing natural selection to act on Y-linked polymorphism are examined. The first incorporates pleiotropic effects of a Y-linked locus, such that viability, segregation distortion, fecundity and sexual selection can all be determined by one locus. It is shown that no set of selection parameters can maintain a stable Y-linked polymorphism. Interaction with the X chromosome is allowed in the next model, with viabilities determined by both X- and Y-linked factors. This model allows four fixation equilibria, two equilibria with X polymorphism and a unique point with both X- and Y-linked polymorphism. Stability analysis shows that the complete polymorphism is never stable. When X- and Y-linked loci influence meiotic drive, it is possible to have all fixation equilibria simultaneously unstable, and yet there is no stable interior equilibrium. Only when viability and meiotic drive are jointly affected by both X- and Y-linked genes can a Y-linked polymorphism be maintained. Unusual dynamics, including stable limit cycles, are generated by this model. Numerical simulations show that only a very small portion of the parameter space admits Y polymorphism, a result that is relevant to the interpretation of levels of Y-DNA sequence variation in natural populations.     

Natural Selection and Age-Structured Populations

This paper studies the properties of a new class of demographic parameters for age-structured populations and analyzes the effect of natural selection on these parameters. Two new demographic variables are introduced: the entropy of a population and the reproductive potential. The entropy of a population measures the variability of the contribution of the different age classes to the stationary population. The reproductive potential measures the mean of the contribution of the different age classes to the Malthusian parameter. The Malthusian parameter is precisely the difference between the entropy and the reproductive potential. The effect of these demographic variables on changes in gene frequency is discussed. The concept of entropy of a genotype is introduced and it is shown that in a random mating population in Hardy-Weinberg equilibrium and under slow selection, the rate of change of entropy is equal to the genetic variance in entropy minus the covariance in entropy and reproductive potential. This result is an information theoretic analog of Fisher''s fundamental theorem of natural selection.     

Selection of a Mixed Culture of Cellulolytic Thermophilic Anaerobes from Various Natural Sources

Microbial associations capable of converting cellulose-containing substrates to ethanol and organic acids were isolated from natural sources. The resulting mixed cultures utilized cellulose, cellobiose, glucose, maize residue, cotton, and flax boon producing ethanol (up to 0.9 g/l) and acetic acid (up to 0.8 g/l). The most complete conversion of cellulose-containing substrates occurred at 60°C and pH 7.0. The selected association of thermophilic anaerobic bacteria produced 0.64 g of ethanol per g substrate utilized at the ethanol/acetate ratio 4.7 : 1.     

Interaction between Natural Selection for Heterozygotes and Directional Selection

Significant correlations between allelic frequencies and environmental variables in a number of insect species have been demonstrated by multivariate techniques. Since many environmental variables show a strong relationship to geographic location and since gene flow between populations can also produce patterns of gene frequencies which are related to the geographic location, both selection and gene-flow hypotheses are consistent with the observed correlations. The genetic variables can be corrected for geographic location and so for linear gene-flow patterns. If, after correction, the genetic variables still show significant correlations with similarly corrected environmental variables, then these correlations are consistent with hypotheses of selection but not of gene flow. The data of Johnson and Schaffer (1973) have been reanalyzed using the method of canonical correlation after correction for geographical location by means of multiple regression. Five of the nine loci studied exhibit significant canonical correlations. These results, under the assumption of linear gene flow, support hypotheses of selective action of environmental variables in the genotype-environment relationships observed.     

Teleosemantics Without Natural Selection

Ruth Millikan and others advocate theories which attempt to naturalize wide mental content (e.g. beliefs’ truth conditions) in terms of function in the teleological sense, where a function is constituted in part by facts concerning past natural selection involving ancestors of a current entity. I argue that it is a mistake to base content on selection. Content should instead be based on functions which though historical, do not involve selection. I sketch an account of such functions.     

Gene Network Polymorphism Is the Raw Material of Natural Selection: The Selfish Gene Network Hypothesis

Population genetics, the mathematical theory of modern evolutionary biology, defines evolution as the alteration of the frequency of distinct gene variants (alleles) differing in fitness over the time. The major problem with this view is that in gene and protein sequences we can find little evidence concerning the molecular basis of phenotypic variance, especially those that would confer adaptive benefit to the bearers. Some novel data, however, suggest that a large amount of genetic variation exists in the regulatory region of genes within populations. In addition, comparison of homologous DNA sequences of various species shows that evolution appears to depend more strongly on gene expression than on the genes themselves. Furthermore, it has been demonstrated in several systems that genes form functional networks, whose products exhibit interrelated expression profiles. Finally, it has been found that regulatory circuits of development behave as evolutionary units. These data demonstrate that our view of evolution calls for a new synthesis. In this article I propose a novel concept, termed the selfish gene network hypothesis, which is based on an overall consideration of the above findings. The major statements of this hypothesis are as follows. (1) Instead of individual genes, gene networks (GNs) are responsible for the determination of traits and behaviors. (2) The primary source of microevolution is the intraspecific polymorphism in GNs and not the allelic variation in either the coding or the regulatory sequences of individual genes. (3) GN polymorphism is generated by the variation in the regulatory regions of the component genes and not by the variance in their coding sequences. (4) Evolution proceeds through continuous restructuring of the composition of GNs rather than fixing of specific alleles or GN variants.     

Natural and Sexual Selection on Many Loci

A method is developed that describes the effects on an arbitrary number of autosomal loci of selection on haploid and diploid stages, of nonrandom mating between haploid individuals, and of recombination. We provide exact recursions for the dynamics of allele frequencies and linkage disequilibria (nonrandom associations of alleles across loci). When selection is weak relative to recombination, our recursions provide simple approximations for the linkage disequilibria among arbitrary combinations of loci. We show how previous models of sex-independent natural selection on diploids, assortative mating between haploids, and sexual selection on haploids can be analyzed in this framework. Using our weak-selection approximations, we derive new results concerning the coevolution of male traits and female preferences under natural and sexual selection. In particular, we provide general expressions for the intensity of linkage-disequilibrium induced selection experienced by loci that contribute to female preferences for specific male traits. Our general results support the previous observation that these indirect selection forces are so weak that they are unlikely to dominate the evolution of preference-producing loci.     

Linkage and the Limits to Natural Selection

The probability of fixation of a favorable mutation is reduced if selection at other loci causes inherited variation in fitness. A general method for calculating the fixation probability of an allele that can find itself in a variety of genetic backgrounds is applied to find the effect of substitutions, fluctuating polymorphisms, and deleterious mutations in a large population. With loose linkage, r, the effects depend on the additive genetic variance in relative fitness, var (W), and act by reducing effective population size by (N/N(e)) = 1 + var (W)/2r(2). However, tightly linked loci can have a substantial effect not predictable from N(e). Linked deleterious mutations reduce the fixation probability of weakly favored alleles by exp(-2U/R), where U is the total mutation rate and R is the map length in Morgans. Substitutions can cause a greater reduction: an allele with advantage s < s(crit) = (π(2)/6) log(e) (S/s)[var(W)/R] is very unlikely to be fixed. (S is the advantage of the substitution impeding fixation.) Fluctuating polymorphisms at many (n) linked loci can also have a substantial effect, reducing fixation probability by exp [ &2Kn var(W)/R] [K = -1/E((u - u)(2)/uv) depending on the frequencies (u,v) at the selected polymorphisms]. Hitchhiking due to all three kinds of selection may substantially impede adaptation that depends on weakly favored alleles.