Evolutionary connectionism: algorithmic principles underlying the evolution of biological organisation in evo-devo, evo-eco and evolutionary transitions

The mechanisms of variation, selection and inheritance, on which evolution by natural selection depends, are not fixed over evolutionary time. Current evolutionary biology is increasingly focussed on understanding how the evolution of developmental organisations modifies the distribution of phenotypic variation, the evolution of ecological relationships modifies the selective environment, and the evolution of reproductive relationships modifies the heritability of the evolutionary unit. The major transitions in evolution, in particular, involve radical changes in developmental, ecological and reproductive organisations that instantiate variation, selection and inheritance at a higher level of biological organisation. However, current evolutionary theory is poorly equipped to describe how these organisations change over evolutionary time and especially how that results in adaptive complexes at successive scales of organisation (the key problem is that evolution is self-referential, i.e. the products of evolution change the parameters of the evolutionary process). Here we first reinterpret the central open questions in these domains from a perspective that emphasises the common underlying themes. We then synthesise the findings from a developing body of work that is building a new theoretical approach to these questions by converting well-understood theory and results from models of cognitive learning. Specifically, connectionist models of memory and learning demonstrate how simple incremental mechanisms, adjusting the relationships between individually-simple components, can produce organisations that exhibit complex system-level behaviours and improve the adaptive capabilities of the system. We use the term “evolutionary connectionism” to recognise that, by functionally equivalent processes, natural selection acting on the relationships within and between evolutionary entities can result in organisations that produce complex system-level behaviours in evolutionary systems and modify the adaptive capabilities of natural selection over time. We review the evidence supporting the functional equivalences between the domains of learning and of evolution, and discuss the potential for this to resolve conceptual problems in our understanding of the evolution of developmental, ecological and reproductive organisations and, in particular, the major evolutionary transitions

Impacts of Parasites in Early Life: Contrasting Effects on Juvenile Growth for Different Family Members

Parasitism experienced early in ontogeny can have a major impact on host growth, development and future fitness, but whether siblings are affected equally by parasitism is poorly understood. In birds, hatching asynchrony induced by hormonal or behavioural mechanisms largely under parental control might predispose young to respond to infection in different ways. Here we show that parasites can have different consequences for offspring depending on their position in the family hierarchy. We experimentally treated European Shag (Phalacrocorax aristoteli) nestlings with the broad-spectrum anti-parasite drug ivermectin and compared their growth rates with nestlings from control broods. Average growth rates measured over the period of linear growth (10 days to 30 days of age) and survival did not differ for nestlings from treated and control broods. However, when considering individuals within broods, parasite treatment reversed the patterns of growth for individual family members: last-hatched nestlings grew significantly slower than their siblings in control nests but grew faster in treated nests. This was at the expense of their earlier-hatched brood-mates, who showed an overall growth rate reduction relative to last-hatched nestlings in treated nests. These results highlight the importance of exploring individual variation in the costs of infection and suggest that parasites could be a key factor modulating within-family dynamics, sibling competition and developmental trajectories from an early age

A Test of Evolutionary Policing Theory with Data from Human Societies

In social groups where relatedness among interacting individuals is low, cooperation can often only be maintained through mechanisms that repress competition among group members. Repression-of-competition mechanisms, such as policing and punishment, seem to be of particular importance in human societies, where cooperative interactions often occur among unrelated individuals. In line with this view, economic games have shown that the ability to punish defectors enforces cooperation among humans. Here, I examine a real-world example of a repression-of-competition system, the police institutions common to modern human societies. Specifically, I test evolutionary policing theory by comparing data on policing effort, per capita crime rate, and similarity (used as a proxy for genetic relatedness) among citizens across the 26 cantons of Switzerland. This comparison revealed full support for all three predictions of evolutionary policing theory. First, when controlling for policing efforts, crime rate correlated negatively with the similarity among citizens. This is in line with the prediction that high similarity results in higher levels of cooperative self-restraint (i.e. lower crime rates) because it aligns the interests of individuals. Second, policing effort correlated negatively with the similarity among citizens, supporting the prediction that more policing is required to enforce cooperation in low-similarity societies, where individuals' interests diverge most. Third, increased policing efforts were associated with reductions in crime rates, indicating that policing indeed enforces cooperation. These analyses strongly indicate that humans respond to cues of their social environment and adjust cheating and policing behaviour as predicted by evolutionary policing theory

Does Sex Speed Up Evolutionary Rate and Increase Biodiversity?

Most empirical and theoretical studies have shown that sex increases the rate of evolution, although evidence of sex constraining genomic and epigenetic variation and slowing down evolution also exists. Faster rates with sex have been attributed to new gene combinations, removal of deleterious mutations, and adaptation to heterogeneous environments. Slower rates with sex have been attributed to removal of major genetic rearrangements, the cost of finding a mate, vulnerability to predation, and exposure to sexually transmitted diseases. Whether sex speeds or slows evolution, the connection between reproductive mode, the evolutionary rate, and species diversity remains largely unexplored. Here we present a spatially explicit model of ecological and evolutionary dynamics based on DNA sequence change to study the connection between mutation, speciation, and the resulting biodiversity in sexual and asexual populations. We show that faster speciation can decrease the abundance of newly formed species and thus decrease long-term biodiversity. In this way, sex can reduce diversity relative to asexual populations, because it leads to a higher rate of production of new species, but with lower abundances. Our results show that reproductive mode and the mechanisms underlying it can alter the link between mutation, evolutionary rate, speciation and biodiversity and we suggest that a high rate of evolution may not be required to yield high biodiversity

Ecological genetics of inbreeding, outbreeding and immunocompetence in Ranid frogs

Using artificial fertilization, I crossed frogs from different populations to evaluate fitness consequences for the offspring from an inbreeding-outbreeding perspective, and to evaluate quantitative genetic effects on immunocompetence against a fungal pathogen (Saprolegnia). Crosses between closely situated populations of different sizes generated contrasting results for the effects of outbreeding on offspring traits between populations and life history stages, emphasizing the importance of epistatic effects and the difficulties of relying on generalizations when making conservation decisions (e.g., regarding translocations). Experimental infection of frog eggs from six populations with Saprolegnia fungus showed a significant family effect on the degree of infection of eggs and embryos, in particular at lower fertilization success and with a significant temperature × population interaction effect. A paternal genetic effect on fungus resistance was found using a half-sib split design. Furthermore, relatively more eggs were infected when fertilized by sperm from the same, in contrast with a different population. However, there was no evidence for a stronger effect in isolated island populations. Although the mechanistic underpinnings remain unknown, these results suggest substantial levels of genetic variation in resistance to Saprolegnia in natural populations within and among populations. We also found that pre-hatching exposure to Saprolegnia dramatically reduced the size at metamorphosis in the absence of further exposure to the fungus, possible as a delayed effect of impaired embryonic development. However, in contrast to some other amphibians, induced hatching in response to Saprolegnia could not be confirmed. In conclusion, the results suggest that frog populations are genetically diverse even at small geographic scale with frequently strong and unpredictable consequences of in- and outbreeding for the response to stressors

Understanding and researching complexity with Qualitative Comparative Analysis: Evaluating transportation infrastructure projects

This article proposes a complexity-informed framework for evaluating transportation infrastructure projects. The article does this through four steps. First, the properties of infrastructure development projects are discussed. This leads to the conclusion that the specific locality or contextualization of a given project is important for explaining the outcome. Hence, there is a need for an ontology and epistemology that addresses the importance of this contextualization. The second step concerns the development of the prerequisites for a methodological framework that follows from this epistemology and ontology. The third step is the assessment of common infrastructure evaluation methods against these prerequisites. This leads to the conclusion that a comparative case-based approach is the most suitable way to study the relationship between context and outcomes in projects. A framework based on Qualitative Comparative Analysis (QCA) is presented in the fourth step. The article concludes with a discussion of the further development of QCA