The evolution of helping and harming on graphs: the return of the inclusive fitness effect

Evolutionary graph theory has been proposed as providing new fundamental rules for the evolution of co‐operation and altruism. But how do these results relate to those of inclusive fitness theory? Here, we carry out a retrospective analysis of the models for the evolution of helping on graphs of Ohtsuki et al. [Nature (2006) 441, 502] and Ohtsuki & Nowak [Proc. R. Soc. Lond. Ser. B Biol. Sci (2006) 273, 2249]. We show that it is possible to translate evolutionary graph theory models into classical kin selection models without disturbing at all the mathematics describing the net effect of selection on helping. Model analysis further demonstrates that costly helping evolves on graphs through limited dispersal and overlapping generations. These two factors are well known to promote relatedness between interacting individuals in spatially structured populations. By allowing more than one individual to live at each node of the graph and by allowing interactions to vary with the distance between nodes, our inclusive fitness model allows us to consider a wider range of biological scenarios leading to the evolution of both helping and harming behaviours on graphs.     

Bacterial ecology, antibiotics and selection for virulence

The evolution of antibiotic resistance in bacteria is well known. Here I describe possible mechanisms by which an increased rate of re-colonization of vertebrate guts by microbes caused by antibiotic use could lead to selection for increased virulence in currently mutualistic or benign microbes. The importance of understanding both the source and the frequency of colonization in such mutualisms is stressed and the possible importance of pseudo-vertical transmission in the evolution of these systems is discussed. A number of areas requiring experimental investigation are identified.     

A field guide for teaching evolution in the social sciences

The theory of evolution by natural selection has begun to revolutionize our understanding of perception, cognition, language, social behavior, and cultural practices. Despite the centrality of evolutionary theory to the social sciences, many students, teachers, and even scientists struggle to understand how natural selection works. Our goal is to provide a field guide for social scientists on teaching evolution, based on research in cognitive psychology, developmental psychology, and education. We synthesize what is known about the psychological obstacles to understanding evolution, methods for assessing evolution understanding, and pedagogical strategies for improving evolution understanding. We review what is known about teaching evolution about nonhuman species and then explore implications of these findings for the teaching of evolution about humans. By leveraging our knowledge of how to teach evolution in general, we hope to motivate and equip social scientists to begin teaching evolution in the context of their own field.     

Symmetry preservation in the evolution of the genetic code

The standard genetic code is found to exhibit an exact symmetry under a finite group of order 4 known in mathematics as the Klein group. The same symmetry is also present in almost all non-standard codes, mitochondrial as well as nuclear. Analysis of the phylogenetic tree for the evolution of the mitochondrial codes reveals that all changes along the main line of evolution preserve this symmetry, with a tendency towards symmetry enhancement. In the side branches of the evolutionary tree, the majority of changes also respect the symmetry. The few exceptional cases where it is broken correspond to reassignments that appear to be unstable or incomplete. Since the Klein group emerges naturally from the symplectic model for the prebiotic evolution that has led to the standard code, we interpret these results as lending support to the hypothesis that this symmetry has been selected during the evolution of the genetic code, not only before but also after establishment of the standard code.     

Measurement techniques for cellular biomechanics in vitro

Living cells and tissues experience mechanical forces in their physiological environments that are known to affect many cellular processes. Also of importance are the mechanical properties of cells, as well as the microforces generated by cellular processes themselves in their microenvironments. The difficulty associated with studying these phenomena in vivo has led to alternatives such as using in vitro models. The need for experimental techniques for investigating cellular biomechanics and mechanobiology in vitro has fueled an evolution in the technology used in these studies. Particularly noteworthy are some of the new biomicroelectromechanical systems (Bio-MEMS) devices and techniques that have been introduced to the field. We describe some of the cellular micromechanical techniques and methods that have been developed for in vitro studies, and provide summaries of the ranges of measured values of various biomechanical quantities. We also briefly address some of our experiences in using these methods and include modifications we have introduced in order to improve them.     

A Graph Theoretical Approach to Study the Organization of the Cortical Networks during Different Mathematical Tasks

The two core systems of mathematical processing (subitizing and retrieval) as well as their functionality are already known and published. In this study we have used graph theory to compare the brain network organization of these two core systems in the cortical layer during difficult calculations. We have examined separately all the EEG frequency bands in healthy young individuals and we found that the network organization at rest, as well as during mathematical tasks has the characteristics of Small World Networks for all the bands, which is the optimum organization required for efficient information processing. The different mathematical stimuli provoked changes in the graph parameters of different frequency bands, especially the low frequency bands. More specific, in Delta band the induced network increases it’s local and global efficiency during the transition from subitizing to retrieval system, while results suggest that difficult mathematics provoke networks with higher cliquish organization due to more specific demands. The network of the Theta band follows the same pattern as before, having high nodal and remote organization during difficult mathematics. Also the spatial distribution of the network’s weights revealed more prominent connections in frontoparietal regions, revealing the working memory load due to the engagement of the retrieval system. The cortical networks of the alpha brainwaves were also more efficient, both locally and globally, during difficult mathematics, while the fact that alpha’s network was more dense on the frontparietal regions as well, reveals the engagement of the retrieval system again. Concluding, this study gives more evidences regarding the interaction of the two core systems, exploiting the produced functional networks of the cerebral cortex, especially for the difficult mathematics.     

Genetics and the evolutionary process

Population genetics was put forward as a mathematical theory between 1918 and 1932 and played a leading part in the rediscovery of the concept of natural selection. As an autonomous science developing Mendel's laws at the population scale and a key element of the Darwinian theory of evolution, its dual status led its practioners to initially overlook some consequences of Mendelism not accounted for by the Darwinian theory, including random drift and the cost of selection. The latter were put forward on purely theoretical grounds in the 1950s, but their importance was acknowledged only when empirical data on protein evolution and enzyme polymorphism (since 1965) and on DNA variation (since 1983) were obtained. The neutralist/selectionist debate that ensued involved disagreement over the scientific method as well as over the mechanisms of molecular evolution. Population genetics has long assumed the existence of natural selection a priori. It has since recentred around the null hypothesis that molecular evolution is neutral. This new approach, applied to sequence comparison and to the study of linkage disequilibrium, is logically more justified, yet empirical observations derived from it paradoxically show the overwhelming importance of selective effects within genomes.     

Evolution’s past and future: an introduction and partial précis to Stephen Jay Gould’s The Structure of Evolutionary Theory

In The Structure of Evolutionary Theory, his last and largest book on evolution, Stephen Jay Gould conceives of the structure of evolutionary theory since Darwin as comprising three major propositions. First, natural selection is the most important direction-giving force in evolution. Second, it operates at the level of the individual organism. Third, selection can be extrapolated smoothly from its actions on individuals in living species throughout geologic time, to produce the gradual divergence of species and adaptations that characterizes the history of life. Challenges to each of these major propositions, according to Gould, can be of three kinds of severity. The most severe challenges, if true, would nullify one of the major propositions entirely, thus destroying the integration of the theory (and perhaps the logic and support of its other propositions). Other, less severe challenges would revise, modify, and expand the content and scope of one or another of the propositions, but not destroy any of them or the theory in toto. Still other, even less severe challenges add to what is known and extend the scope and possibilities of the theory, but do not call for a revision in its fundamental structure. Gould acknowledges that the theory has withstood all presumptive challenges that would destroy it, and has accommodated those that simply extend and add to it. His principal concern is with those challenges that would revise the theory substantially: for example, if processes other than natural selection were of great importance in evolution; if selection acted in important ways at the level of species and clades, and (or) at the level of the genes, alleles, and chromosomes; and if the extrapolation of what is known from living populations could not by itself explain many patterns of large-scale evolution seen in the fossil record. He thinks that, both through the history of evolution since Darwin and in the present day, challenges that substantially revise these basic propositions are valid, and that the theory needs to integrate them in order to retain the explanatory power that it has had for many decades. To cite this article: K. Padian, C. R. Palevol 2 (2003).     

MATHEMATICS OF KIN‐ AND GROUP‐SELECTION: FORMALLY EQUIVALENT?

Evolutionary game theory is a general mathematical framework that describes the evolution of social traits. This framework forms the basis of many multilevel selection models and is also frequently used to model evolutionary dynamics on networks. Kin selection, which was initially restricted to describe social interactions between relatives, has also led to a broader mathematical approach, inclusive fitness, that can not only describe social evolution among relatives, but also in group structured populations or on social networks. It turns out that the underlying mathematics of game theory is fundamentally different from the approach of inclusive fitness. Thus, both approaches—evolutionary game theory and inclusive fitness—can be helpful to understand the evolution of social traits in group structured or spatially extended populations.     

Alpheus Spring Packard and cave fauna in the evolution debate

Conclusion Packard attempted to incorporate cave fauna into a general theory of evolution that would be consistent with the principle of recapitulation, and would have as the primary mechanism the inheritance of the effects of the environment. Beyond this, he also attempted to demonstrate that the evolution of cave fauna was consistent with progressive evolution. The use he made of comparative anatomy and embryology places him within the tradition of classical morphology that was dominant through much of the last half of the nineteenth century, but of waning importance by the time of Packard's death in 1905. The importance Packard gave to cave fauna as evidence for Lamarckian evolution stimulated interest in the phenomenon; this interest, and references to cave fauna in the scientific literature, declined after his death. Since then, the importance of cave fauna in evolutionary theory has declined from their status as the star evidence in Packard's theory to their present status as a difficult anomaly within the modern synthetic theory.     

Evolutionary biotechnology – theory,facts and perspectives

Molecular evolution has recently been applied in biotechnology which consist of the development of evolutionary strategies in the design of biopolymers with predefined properties and functions. At the heart of this new technology are the in vitro replication and random synthesis of RNA or DNA molecules, producing large libraries of genotypes that are subjected to selection techniques following DARWIN's principle. By means of these evolutionary methods, RNA molecules were derived which specifically bind to predefined target molecules. Ribozymes with new catalytic functions were obtained as well as RNA molecules that are resistant to cleavage by specific RNases. In addition, the catalytic specificities of group I introns, a special class of ribozymes, were modified by variation and selection. Efficient applications of molecular evolution to problems in biotechnology require a fundamental and detailed understanding of the evolutionary process. Two basic questions are of primary importance: (i) How can evolutionary methods be successful as the numbers of possible genotypes are so large that the chance of obtaining a particular sequence by random processes is practically zero, and (ii) how can populations avoid being caught in evolutionary traps corresponding to local fitness optima? This review is therefore concerned with an abridged account of the theory of molecular evolution, as well as its application to biotechnology. We add a brief discussion of new techniques for the massively parallel handling and screening of very small probes as is required for the spatial separation and selection of genotypes. Finally, some imminent prospects concerning the evolutionary design of biopolymers are presented.     

Absolute fitness, relative fitness, and utility

It is well known that (1) natural selection typically favors an allele with both a large mean fitness and a small variance in fitness; and (2) investors typically prefer a portfolio with both a large mean return and a small variance in returns. In the case of investors, this mean-variance trade-off reflects risk aversion; in the case of evolution, the mathematics is straightforward but the result is harder to intuit. In particular, it is harder to understand where, in the mathematics of natural selection, risk aversion arises. Here I present a result that suggests a simple answer to this question. Although my answer is essentially identical to one offered previously, my path to it differs somewhat from previous approaches. Some may find this new approach easier to intuit.     

Molecular evolution and selection pressure in alpha-class carbonic anhydrase family members

Carbonic anhydrases (CA) are ubiquitous, and their involvement in diseases such as hypertension, diabetes, and glaucoma is well known. Most members of this family of metalloenzymes convert carbon dioxide to bicarbonate with the help of a Zn(2+) cofactor. While the expression patterns and kinetic activities of many of these isozymes have been studied, little is known about the differences in the conservation patterns of individual residues. To better understand the molecular evolution of the CA gene family, we created multiple sequence alignments and analyzed the selection pressure (dN/dS ratios) on surface and active site residues in 248 mammalian sequences of the 14 known family members. Using the values found for amino acids of known functional importance (i.e. the three histidines that bind the zinc cofactor) as our baseline, we were able to identify other regions of possible structural and functional importance.     

The Importance of Understanding the Nature of Science for Accepting Evolution

Many students reject evolutionary theory, whether or not they adequately understand basic evolutionary concepts. We explore the hypothesis that accepting evolution is related to understanding the nature of science. In particular, students may be more likely to accept evolution if they understand that a scientific theory is provisional but reliable, that scientists employ diverse methods for testing scientific claims, and that relating data to theory can require inference and interpretation. In a study with university undergraduates, we find that accepting evolution is significantly correlated with understanding the nature of science, even when controlling for the effects of general interest in science and past science education. These results highlight the importance of understanding the nature of science for accepting evolution. We conclude with a discussion of key characteristics of science that challenge a simple portrayal of the scientific method and that we believe should be emphasized in classrooms.     

Conditionally Neutral Phylogenetic Markers of Major Taxa: A New Aspect of the Evolution of Macromolecules

The current phase of molecular phylogenetics can be named the 18S rRNA gene era, which is now approaching the end. To date, almost all phyla of metazoans and many taxa of protists are represented in databases of 18S rRNA gene sequences. The elements of the phylogenetic tree of Metazoa inferred from 18S rRNA genes are characterized by unequal validity: some of them seem to be well grounded; others are not adequately supported, and probably will be revised later. The validity of phylogenetic reconstruction is influenced by two main factors: (1) erroneous grouping of long branches that occur because of abnormally high evolution rate; (2) deficit of phylogenetically informative characters. A method for overcoming these difficulties is suggested in addition to known tools: using phylogenetic markers that are stable within individual taxa and evolve by punctuated equilibrium. These markers are least influenced by the convergence caused by a high evolution rate of the entire gene. The nature of these markers of ancient taxa, paradoxical from the perspective of neutral evolution, is discussed, as well as their importance for establishing monophyly of both new large-scale taxonomic groups of invertebrates (Bilateria + Rhombozoa + Orthonectida + Myxozoa + Cnidaria + Placozoa and Echinodermata + Hemichordata) and some major taxa of Nematoda.     

Conditionally neutral phylogenetic markers of major taxa: a new aspect of the evolution of macromolecules

The current phase of molecular phylogenetics can be named the 18S rRNA gene era, which is now approaching the end. To date, almost all phyla of metazoans and many taxa of protists are represented in databases of 18S rRNA gene sequences. The elements of the phylogenetic tree of Metazoa inferred from 18S rRNA genes are characterized by unequal validity: some of them seem to be well grounded; others are not adequately supported, and probably will be revised later. The validity of phylogenetic reconstruction is influenced by two main factors: (1) erroneous grouping of long branches that occur because of abnormally high evolution rate; (2) deficit of phylogenetically informative characters. A method for overcoming these difficulties is suggested in addition to known tools: using phylogenetic markers that are stable within individual taxa and evolve by punctuated equilibrium. These markers are least influenced by the convergence caused by a high evolution rate of the entire gene. The nature of these markers of ancient taxa, paradoxical from the perspective of neutral evolution, is discussed, as well as their importance for establishing monophyly of both new large-scale taxonomic groups of invertebrates (Bilateria + Rhombozoa + Orthonectida + Myxozoa + Cnidaria + Placozoa and Echinodermata + Hemichordata) and some major taxa of Nematoda.     

The combinatorial distance geometry method for the calculation of molecular conformation. I. A new approach to an old problem

A new approach to the long-standing local minimum problem of molecular energy minimization is proposed. The approach relies upon a field of computer mathematics known as combinatorial optimization, together with methods of conformational analysis derived from distance geometry. The advantages over the usual numerical techniques of optimization are, first, that the algorithms derived are globally convergent, and second, that the mathematical problems involved are well-posed and suitable for study within the modern theory of computational complexity. In this paper we introduce the approach, and describe a computer program based on it.     

Group selection and kin selection: formally equivalent approaches

Inclusive fitness theory, summarised in Hamilton's rule, is a dominant explanation for the evolution of social behaviour. A parallel thread of evolutionary theory holds that selection between groups is also a candidate explanation for social evolution. The mathematical equivalence of these two approaches has long been known. Several recent papers, however, have objected that inclusive fitness theory is unable to deal with strong selection or with non-additive fitness effects, and concluded that the group selection framework is more general, or even that the two are not equivalent after all. Yet, these same problems have already been identified and resolved in the literature. Here, I survey these contemporary objections, and examine them in the light of current understanding of inclusive fitness theory.     

Chance as an explanatory factor in evolutionary biology.

Darwinian evolutionary biology has often been criticized for appealing to the notion of 'chance' in its explanations. According to some critics, such appeals exhibit the explanatory poverty of evolutionary theory. In response, defenders of Darwinism sometimes downplay the importance of 'chance' in evolution. I believe that both of these approaches are mistaken. The main thesis of this paper is that the term 'chance' encompasses a number of distinct concepts, and that at least some of these concepts serve essential explanatory functions in evolutionary biology. This claim is defended by way of an historical survey of the major concepts of 'chance' in the history of evolutionary biology, especially the concepts used by Jean Baptiste Lamarck, Charles Darwin, and Sewall Wright. An examination of their biologies shows how the concepts of 'chance' used cohere with their major scientific objectives and methods. These concepts survive and continue to function as important explanatory factors in contemporary evolutionary biology. Examples of such usage are given, and the explanatory status of 'chance' assessed.     

Caspar carboxylates: the structural basis of tobamovirus disassembly.

Carboxylate groups have been known for many years to drive the disassembly of simple viruses, including tobacco mosaic virus (TMV). The identities of the carboxylate groups involved and the mechanism by which they initiate disassembly have not, however, been clear. Structures have been determined at resolutions between 2.9 and 3.5 A for five tobamoviruses by fiber diffraction methods. Site-directed mutagenesis has also been used to change numerous carboxylate side chains in TMV to the corresponding amides. Comparison of the stabilities of the various mutant viruses shows that disassembly is driven by a much more complex set of carboxylate interactions than had previously been postulated. Despite the importance of the carboxylate interactions, they are not conserved during viral evolution. Instead, it appears that during evolution, patches of electrostatic interaction drift across viral subunit interfaces. The flexibility of these interactions confers a considerable advantage on the virus, enabling it to change its surface structure rapidly and thus evade host defenses.