Multilevel selection theory and evidence: a critique of Gardner, 2015.

Gardner (2015) recently developed a model of a 'Genetical Theory of Multilevel Selection, which is a thoughtfully developed, but flawed model. The model's flaws appear to be symptomatic of common misunderstandings of the multi level selection (MLS) literature and the recent quantitative genetic literature. I use Gardner's model as a guide for highlighting how the MLS literature can address the misconceptions found in his model, and the kin selection literature in general. I discuss research on the efficacy of group selection, the roll of indirect genetic effects in affecting the response to selection and the heritability of group-level traits. I also discuss why the Price multilevel partition should not be used to partition MLS, and why contextual analysis and, by association, direct fitness are appropriate for partitioning MLS. Finally, I discuss conceptual issues around questions concerning the level at which fitness is measured, the units of selection, and I present a brief outline of a model of selection in class-structured populations. I argue that the results derived from the MLS research tradition can inform kin selection research and models, and provide insights that will allow researchers to avoid conceptual flaws such as those seen in the Gardner model.

Quantitative trait loci and gene interaction: the quantitative genetics of metapopulations.

Genetic population differentiation is typically viewed as differentiation of population means. However, several theories of evolution and speciation postulate that populations differentiate not only with respect to the population means, but also with respect to the effects of alleles within these populations. I develop herein a measure of population differentiation for the 'local average effects' of alleles, where local average effect is defined as the average effect of an allele in a deme measured as a deviation from the metapopulation mean. The differentiation for local average effects has two components, a component attributable to the population mean and a residual component that is attributable to changes in the local average effects independent of the population mean. The variance in local average effects attributable to the population mean is measured as the variance in the mean local average effect of all alleles. The variance in the residual local average effects is measured as the difference between the variance local average effects of individual alleles and the variance in the mean local average effects of all alleles. Differentiation for population means and differentiation for residual local average effects need not be related. I show that when there is only additive gene action, populations can differentiate for population means, but not for residual local average effects. However, if there is gene interaction then populations can also differentiate for local average effects of alleles. The consequence of this differentiation is that the local average effects of alleles change relative to each other such that an allele that is favoured by selection in one population may be removed by selection in other populations. I discuss the evolutionary consequences of differentiation for local average effects, and the interpretation of QTL data in light of this model.

Experimental studies of group selection: what do they tell us about group selection in nature?

The study of group selection has developed along two autonomous lines. One approach, which we refer to as the adaptationist school, seeks to understand the evolution of existing traits by examining plausible mechanisms for their evolution and persistence. The other approach, which we refer to as the genetic school, seeks to examine how currently acting artificial or natural selection changes traits within populations and focuses on current evolutionary change. The levels of selection debate lies mainly within the adaptationist school, whereas the experimental studies of group selection lie within the genetic school. Because of the very different traditions and goals of these two schools, the experimental studies of group selection have not had a major impact on the group selection debate. We review the experimental results of the genetic school in the context of the group selection controversy and address the following questions: Under what conditions is group selection effective? What is the genetic basis of a response to group selection? How common is group selection in nature?

Wright's shifting balance theory: an experimental study.

Experimental confirmation of Wright's shifting balance theory of evolution, one of the most comprehensive theories of adaptive evolution, is presented. The theory is regarded by many as a cornerstone of modern evolutionary thought, but there has been little direct empirical evidence supporting it. Some of its underlying assumptions are viewed as contradictory, and the existence and efficacy of the theory's fundamental adaptive process, interdemic selection, is the focus of controversy. Interdemic selection was imposed on large arrays of laboratory populations of the flour beetle Tribolium castaneum in the manner described by Wright: the differential dispersion of individuals from demes of high fitness into demes of low fitness. A significant increase in average fitness was observed in the experimental arrays when compared to control populations with equivalent but random migration rates. The response was not proportional to the selection differential: The largest response occurred with interdemic selection every two generations rather than every generation or every three generations. The results indicate that the interdemic phase of Wright's shifting balance theory can increase average fitness and suggest that gene interactions are involved in the observed response.

Neurosecretory cells: daily rhythmicity in Leiobunum longipes.

A fluorometric-analysis procedure, used to quantitate indoles, confirmed the presence of 5-hydroxytryptamine (5-HT) in an arachnid; there was unimodal cyclic production of 5-HT in brain and intestinal tissues over a 24-hour period. The same tissues produced 5-HT after 80-day culture; bimodal cyclic output was indicated during continuous 24-hour study. One peak occurred at 0200 hours, at the same time as the peak in vivo, suggesting an endogenously controlled mechanism of secretion. The second peak occurred at midmorning, a time when production in vivo was lowest, suggesting that there is a possible feedback-control mechanism in the organism that inhibits the endogenous output of 5-HT.