The long‐running debate about the role of selection in maintaining genetic variation has been given new impetus by the discovery of hundreds of seasonally oscillating polymorphisms in wild Drosophila, possibly stabilized by an alternating summer‐winter selection regime. Historically, there has been skepticism about the potential of temporal variation to balance polymorphism, because selection must be strong to have a meaningful stabilizing effect—unless dominance also varies over time (“reversal of dominance”). Here, we develop a simplified model of seasonally variable selection that simultaneously incorporates four different stabilizing mechanisms, including two genetic mechanisms (“cumulative overdominance” and reversal of dominance), as well as ecological “storage” (“protection from selection” and boom‐bust demography). We use our model to compare the stabilizing effects of these mechanisms. Although reversal of dominance has by far the greatest stabilizing effect, we argue that the three other mechanisms could also stabilize polymorphism under plausible conditions, particularly when all three are present. With many loci subject to diminishing returns epistasis, reversal of dominance stabilizes many alleles of small effect. This makes the combination of the other three mechanisms, which are incapable of stabilizing small effect alleles, a better candidate for stabilizing the detectable frequency oscillations of large effect alleles.

To detect a direction to evolution, without the pitfalls of reconstructing ancestral states, we need to compare "more evolved" to "less evolved" entities. But because all extant species have the same common ancestor, none are chronologically more evolved than any other. However, different gene families were born at different times, allowing us to compare young protein-coding genes to those that are older and hence have been evolving for longer. To be retained during evolution, a protein must not only have a function, but must also avoid toxic dysfunction such as protein aggregation. There is conflict between the two requirements: hydrophobic amino acids form the cores of protein folds, but also promote aggregation. Young genes avoid strongly hydrophobic amino acids, which is presumably the simplest solution to the aggregation problem. Here we show that young genes' few hydrophobic residues are clustered near one another along the primary sequence, presumably to assist folding. The higher aggregation risk created by the higher hydrophobicity of older genes is counteracted by more subtle effects in the ordering of the amino acids, including a reduction in the clustering of hydrophobic residues until they eventually become more interspersed than if distributed randomly. This interspersion has previously been reported to be a general property of proteins, but here we find that it is restricted to old genes. Quantitatively, the index of dispersion delineates a gradual trend, i.e., a decrease in the clustering of hydrophobic amino acids over billions of years.

The extended evolutionary synthesis invokes a role for development in shaping adaptive evolution, which in population genetics terms corresponds to mutation-biased adaptation. Critics have claimed that clonal interference makes mutation-biased adaptation rare. We consider the behaviour of two simultaneously adapting traits, one with larger mutation rate U , the other with larger selection coefficient s , using asexual travelling wave models. We find that adaptation is dominated by whichever trait has the faster rate of adaptation v in isolation, with the other trait subject to evolutionary stalling. Reviewing empirical claims for mutation-biased adaptation, we find that not all occur in the ‘origin-fixation’ regime of population genetics where v is only twice as sensitive to s as to U . In some cases, differences in U are at least ten to twelve times larger than differences in s , as needed to cause mutation-biased adaptation even in the ‘multiple mutations’ regime. Surprisingly, when U > s in the ‘diffusive-mutation’ regime, the required sensitivity ratio is also only two, despite pervasive clonal interference. Given two traits with identical v , the benefit of having higher s is surprisingly small, occurring largely when one trait is at the boundary between the origin-fixation and multiple mutations regimes.

A part analytical, part numerical ideal MHD analysis of low-frequency Alfvén wave physics in the H-1 stellarator is given. The three-dimensional, compressible ideal spectrum for H-1 is presented and it is found that despite the low β of H-1 plasmas (β ≈ 10 −4 ), significant Alfvén-acoustic interactions occur at low frequencies.Several quasi-discrete modes are found with the three-dimensional linearised ideal MHD eigenmode solver CAS3D, including beta-induced Alfvén eigenmode (BAE)type modes in beta-induced gaps. The strongly shaped, low-aspect ratio magnetic geometry of H-1 causes CAS3D convergence difficulties requiring the inclusion of many Fourier harmonics for the parallel component of the fluid displacement eigenvector even for shear wave motions. The highest beta-induced gap reproduces large parts of the observed configurational frequency dependencies in the presence of hollow temperature profiles. *

Abstract. Tree cover varies enormously across tropical ecosystems-from arid savannas to closed rain forests-and yet a general predictive theory of tropical tree cover remains elusive. Here we use the maximum-entropy method to predict the most likely sample frequency distribution of ecosystems with different tree and grass fractional cover if balance between water supply and demand were the dominant constraint on community assembly. Assuming a hierarchy of individual plant water demand in which trees require more water than grasses, we reproduce observed trends in the means and the upper and lower limits of tropical tree and grass cover across the entire spectrum of tropical ecosystem water supply. Finer details not captured by our predictions indicate the influence of additional factors, such as disturbance. Our results challenge the view that tropical tree-grass coexistence is largely sustained by disturbances in moist environments (''unstable'' coexistence) with water supply playing a dominant role only in arid conditions (''stable'' coexistence). More generally, they suggest that macroecological patterns can be understood and predicted as the most likely outcome of a large number of stochastic processes being played out within a relatively small number of ecological constraints.