The Hamiltonian view of social evolution

Hamilton’s Rule, named after the evolutionary biologist Bill Hamilton, and the related concepts of inclusive fitness and kin selection, have been the bedrock of the study of social evolution for the past half century. In ’The Philosophy of Social Evolution’ (Oxford University Press, 2017), Jonathan Birch provides a comprehensive introduction to the conceptual foundations of the Hamiltonian view of social evolution, and a passionate defence of its enduring value in face of the recent high profile criticism. In this review essay, I first outline the version of Hamilton’s Rule defended by Birch, dubbed Hamilton’s Rule General, and its derivation. With this in place, I then navigate through the fierce disagreements that Hamilton’s Rule generates among social evolution researchers and evaluate Birch’s central argument of the book that HRG serves as an organizing framework for social evolution research under which we can compare and interpret more detailed causal models. I then spend the remainder of the review discussing what I take to be three of the most exciting implications of Hamilton’s thinking raised by Birch: (1) the extension of Hamilton’s Rule to mobile genetic elements, (2) maximization of inclusive fitness models and the idea of adaptation as organism design, and (3) the relationship between Hamilton’s approaches to social behaviour and the gene’s-eye view of evolution

The Hamiltonian view of social evolution

Hamilton’s Rule, named after the evolutionary biologist Bill Hamilton, and the related concepts of inclusive fitness and kin selection, have been the bedrock of the study of social evolution for the past half century. In ’The Philosophy of Social Evolution’ (Oxford University Press, 2017), Jonathan Birch provides a comprehensive introduction to the conceptual foundations of the Hamiltonian view of social evolution, and a passionate defence of its enduring value in face of the recent high profile criticism. In this review essay, I first outline the version of Hamilton’s Rule defended by Birch, dubbed Hamilton’s Rule General, and its derivation. With this in place, I then navigate through the fierce disagreements that Hamilton’s Rule generates among social evolution researchers and evaluate Birch’s central argument of the book that HRG serves as an organizing framework for social evolution research under which we can compare and interpret more detailed causal models. I then spend the remainder of the review discussing what I take to be three of the most exciting implications of Hamilton’s thinking raised by Birch: (1) the extension of Hamilton’s Rule to mobile genetic elements, (2) maximization of inclusive fitness models and the idea of adaptation as organism design, and (3) the relationship between Hamilton’s approaches to social behaviour and the gene’s-eye view of evolution

Enforcement is central to the evolution of cooperation.

Cooperation occurs at all levels of life, from genomes, complex cells and multicellular organisms to societies and mutualisms between species. A major question for evolutionary biology is what these diverse systems have in common. Here, we review the full breadth of cooperative systems and find that they frequently rely on enforcement mechanisms that suppress selfish behaviour. We discuss many examples, including the suppression of transposable elements, uniparental inheritance of mitochondria and plastids, anti-cancer mechanisms, reciprocation and punishment in humans and other vertebrates, policing in eusocial insects and partner choice in mutualisms between species. To address a lack of accompanying theory, we develop a series of evolutionary models that show that the enforcement of cooperation is widely predicted. We argue that enforcement is an underappreciated, and often critical, ingredient for cooperation across all scales of biological organization

Exogenous selection shapes germination behaviour and seedling traits of populations at different altitudes in a Senecio hybrid zone

Background and Aims The Senecio hybrid zone on Mt Etna, Sicily, is characterized by steep altitudinal clines in quantitative traits and genetic variation. Such clines are thought to be maintained by a combination of ‘endogenous' selection arising from genetic incompatibilities and environment-dependent ‘exogenous' selection leading to local adaptation. Here, the hypothesis was tested that local adaptation to the altitudinal temperature gradient contributes to maintaining divergence between the parental species, S. chrysanthemifolius and S. aethnensis. Methods Intra- and inter-population crosses were performed between five populations from across the hybrid zone and the germination and early seedling growth of the progeny were assessed. Key Results Seedlings from higher-altitude populations germinated better under low temperatures (9-13 °C) than those from lower altitude populations. Seedlings from higher-altitude populations had lower survival rates under warm conditions (25/15 °C) than those from lower altitude populations, but also attained greater biomass. There was no altitudinal variation in growth or survival under cold conditions (15/5 °C). Population-level plasticity increased with altitude. Germination, growth and survival of natural hybrids and experimentally generated F1s generally exceeded the worse-performing parent. Conclusions Limited evidence was found for endogenous selection against hybrids but relatively clear evidence was found for divergence in seed and seedling traits, which is probably adaptive. The combination of low-temperature germination and faster growth in warm conditions might enable high-altitude S. aethnensis to maximize its growth during a shorter growing season, while the slower growth of S. chrysanthemifolius may be an adaptation to drought stress at low altitudes. This study indicates that temperature gradients are likely to be an important environmental factor generating and maintaining adaptive divergence across the Senecio hybrid zone on Mt Etn

Genes and organisms in the legacy of the modern synthesis

The gene's-eye view of evolution is an influential but contentious perspective on biology. It emerged in the aftermath of the Modern Synthesis and both proponents and detractors have stressed the link between the two. In particular, both the Modern Synthesis and the gene's-eye view have been criticized for overemphasizing the role genes at the expense of organisms in evolutionary explanations. In this chapter, I discuss the connection between the Modern Synthesis and the gene’s-eye view and evaluate the status of genes and organisms in contemporary biology. I show that while the gene’s-eye view traces its origin back to the Modern Synthesis, it can most accurately be said to represent a specific – adaptationist and gene-centric – version of it. To assess the role of genes and organisms, I examine the intimate relationship between the gene’s-eye view and another post-Synthesis development, the concept of inclusive fitness. I argue that the popularity and influence of inclusive fitness theory demonstrate that the individual organism remains safe at the heart of modern evolutionary biology

The electronic structure of poly(pyridine-2,5-diyl) investigated by soft x-ray absorption and emission spectroscopies

The electronic structure of the poly-pyridine conjugated polymer has been investigated by resonant and nonresonant inelastic X-ray scattering and X-ray absorption spectroscopies using synchrotron radiation. The measurements were made for both the carbon and nitrogen contents of the polymer. The analysis of the spectra has been carried out in comparison with molecular orbital calculations taking the repeat-unit cell as a model molecule of the polymer chain. The simulations indicate no significant differences in the absorption and in the non-resonant X-ray scattering spectra for the different isomeric geometries, while some isomeric dependence of the resonant spectra is predicted. The resonant emission spectra show depletion of the {\pi} electron bands in line with symmetry selection and momentum conservation rules. The effect is most vizual for the carbon spectra; the nitrogen spectra are dominated by lone pair n orbital emission of {\sigma} symmetry and are less frequency dependent.Comment: 11 pages, 7 figures, 1 table, http://www.sciencedirect.com/science/article/pii/S030101049800262

Competition between decay and dissociation of core-excited OCS studied by X-ray scattering

We show the first evidence of dissociation during resonant inelastic soft X-ray scattering. Carbon and oxygen K-shell and sulfur L-shell resonant and non-resonant X-ray emission spectra were measured using monochromatic synchrotron radiation for excitation and ionization. After sulfur, L2,3 -> {\pi}*, {\sigma}* excitation, atomic lines are observed in the emission spectra as a consequence of competition between de-excitation and dissociation. In contrast the carbon and oxygen spectra show weaker line shape variations and no atomic lines. The spectra are compared to results from ab initio calculations and the discussion of the dissociation paths is based on calculated potential energy surfaces and atomic transition energies.Comment: 12 pages, 6 pictures, 2 tables, http://link.aps.org/doi/10.1103/PhysRevA.59.428

Solar cycle modulation of Titan's ionosphere

This is the publisher's version, also available electronically from http://onlinelibrary.wiley.com/doi/10.1002/jgra.50463/abstractDuring the six Cassini Titan flybys T83–T88 (May 2012 to November 2012) the electron density in the ionospheric peak region, as measured by the radio and plasma wave science instrument/Langmuir probe, has increased significantly, by 15–30%, compared to previous average. These measurements suggest that a long‒term change has occurred in the ionosphere of Titan, likely caused by the rise to the new solar maximum with increased EUV fluxes. We compare measurements from TA, TB, and T5, from the declining phase of solar cycle 23 to the recent T83–T88 measurements during cycle 24, since the solar irradiances from those two intervals are comparable. The peak electron densities normalized to a common solar zenith angle Nnorm from those two groups of flybys are comparable but increased compared to the solar minimum flybys (T16–T71). The integrated solar irradiance over the wavelengths 1–80nm, i.e., the solar energy flux, Fe, correlates well with the observed ionospheric peak density values. Chapman layer theory predicts that inline image, with k=0.5. We find observationally that the exponent k=0.54±0.18. Hence, the observations are in good agreement with theory despite the fact that many assumptions in Chapman theory are violated. This is also in good agreement with a similar study by Girazian and Withers (2013) on the ionosphere of Mars. We use this power law to estimate the peak electron density at the subsolar point of Titan during solar maximum conditions and find it to be about 6500cm−3, i.e., 85–160% more than has been measured during the entire Cassini mission