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Direct Reciprocity Under Uncertainty Does Not Explain One-Shot Cooperation, But It Can Explain Norm Psychology

Humans in many societies cooperate in economic experiments at much higher levels than would be expected if their goal was maximizing economic returns even when interactions are anonymous and one-shot. This is a puzzle because paying a cost to benefit another player in one-shot interactions has no direct benefit to the cooperator. This paper explores the logic of two competing evolutionary hypotheses to explain this behavior. The \"norm psychology\" hypothesis holds that a players choice of strategy is heavily influenced by socially-learned cultural norms. Its premise is that over the course of human evolutionary history, cultural norms varied considerably across human societies and through a process of gene-culture co-evolution, humans evolved mechanisms to learn and adopt the norms of their particular society. The \"evolutionary mismatch\" hypothesis holds that pro-social preferences evolved genetically in our hunter-gatherer past where one-shot anonymous interactions were rare and these evolved \"protocols\" for cooperation are misapplied in modern, laboratory, conditions. I compare these hypotheses by adopting a well-known model of the mismatch hypothesis. I show that the cooperation generated by the model is based on a flawed assumption - that the best thing to do is cooperate in a repeated game. I show that repeated games generate a great diversity of behavioral equilibria, in support of the norm psychology hypothesiss premise. When interaction is repeated, adopting local norms is a more evolutionarily successful strategy than automatically cooperating. If various groups are at different behavioral equilibria, then cultural selection between groups tends to select for cooperative behavior.

Evolutionary Biology

Revisiting the effect of population size on cumulative cultural evolution

Henrich (2004) argued that larger populations can better maintain complex technologies because they contain more highly skilled people whom others can imitate. His original model, however, did not distinguish the effects of population size from population density or network size; a learners social network included the entire population. Does population size remain important when populations are subdivided and networks are realistically small? I use a mathematical model to show that population size has little effect on equilibrium levels of mean skill under a wide range of conditions. The effects of network size and transmission error rate usually overshadow that of population size. Population size can, however, affect the rate at which a population approaches equilibrium, by increasing the rate at which innovations arise. This effect is small unless innovation is very rare. Whether population size predicts technological complexity in the real world, then, depends on whether technological evolution is innovation-limited and short of equilibrium. The effect of population \"connectedness,\" via migration or trade, is similar. I discuss the results of this analysis in light of the current empirical debate.

Evolutionary Biology

Ecological and Evolutionary Oscillations in Host-Parasite Population Dynamics, and The Red Queen

In a host-parasite system, the constitutive interaction among the species, regulated by the growth rates and functional response, may induce populations to approach equilibrium or sometimes to exhibit simple cycles or peculiar oscillations, such as chaos. A large carrying capacity coupled with appropriate parasitism effectiveness frequently drives long-term apparent oscillatory dynamics in population size. We name these oscillations due to the structure of the constitutive interaction among species as ecological.\n\nOn the other hand, there are also exceptional cases when the evolving quantitative traits of the hosts and parasites induce oscillating population size, which we call as evolutionary. This oscillatory behavior is dependent on the speed of evolutionary adaptation and degree of evolutionary trade-off. A moderate level of negative trade-off is essential for the existence of oscillations. Evolutionary oscillations due to the host-parasite coevolution (known as the Red Queen) can be observed beyond the ecological oscillations, especially when there are more than two competing species involved.\n\nOne Sentence SummaryWe investigate several cases yielding to oscillating host-parasite populations, and we found that the Red Queen hypothesis can explain some of the exceptional cases.\n\nGraphical Abstract:\n\nO_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=114 SRC=\"FIGDIR/small/001735_ufig1.gif\" ALT=\"Figure 1\">\nView larger version (28K):\norg.highwire.dtl.DTLVardef@1280478org.highwire.dtl.DTLVardef@3d6e28org.highwire.dtl.DTLVardef@10b2718org.highwire.dtl.DTLVardef@133a78d_HPS_FORMAT_FIGEXP M_FIG C_FIG

Evolutionary Biology

Population genomics of Saccharomyces cerevisiae human isolates: passengers, colonizers, invaders.

The quest for the ecological niches of Saccharomyces cerevisiae ranged from wineries to oaks and more recently to the gut of Crabro Wasps. Here we propose the role of the human gut in shaping S. cerevisiae evolution, presenting the genetic structure of a previously unknown population of yeasts, associated with Crohns disease, providing evidence for clonal expansion within humans gut. To understand the role of immune function in the human-yeast interaction we classified strains according to their immunomodulatory properties, discovering a set of genetically homogeneous isolates, capable of inducing anti-inflammatory signals via regulatory T cells proliferation, and on the contrary, a positive association between strain mosaicism and ability to elicit inflammatory, IL-17 driven, immune responses. The approach integrating genomics with immune phenotyping showed selection on genes involved in sporulation and cell wall remodeling as central for the evolution of S. cerevisiae Crohns strains from passengers to commensals to potential pathogens.

Evolutionary Biology

Holsteins Favor Heifers, Not Bulls: Biased Milk Production Programmed during Pregnancy as a Function of Fetal Sex

Mammalian females pay high energetic costs for reproduction, the greatest of which is imposed by lactation. The synthesis of milk requires, in part, the mobilization of bodily reserves to nourish developing young. Numerous hypotheses have been advanced to predict how mothers will differentially invest in sons and daughters, however few studies have addressed sex-biased milk synthesis. Here we leverage the dairy cow model to investigate such phenomena. Using 2.39 million lactation records from 1.49 million dairy cows, we demonstrate that the sex of the fetus influences the capacity of the mammary gland to synthesize milk during lactation. Cows favor daughters, producing significantly more milk for daughters than for sons across lactation. Using a sub-sample of this dataset (N = 113,750 subjects) we further demonstrate that the effects of fetal sex interact dynamically across parities, whereby the sex of the fetus being gestated can enhance or diminish the production of milk during an established lactation. Moreover the sex of the fetus gestated on the first parity has persistent consequences for milk synthesis on the subsequent parity. Specifically, gestation of a daughter on the first parity increases milk production by [~]445 kg over the first two lactations. Our results identify a dramatic and sustained programming of mammary function by offspring in utero. Nutritional and endocrine conditions in utero are known to have pronounced and long-term effects on progeny, but the ways in which the progeny has sustained physiological effects on the dam have received little attention to date.

Evolutionary Biology

Evolutionary dynamics of shared niche construction

Many species engage in niche construction that ultimately leads to an increase in the carrying capacity of the population. We have investigated how the specificity of this behaviour affects evolutionary dynamics using a set of coupled logistic equations, where the carrying capacity of each genotype consists of two components: an intrinsic part and a contribution from all genotypes present in the population. The relative contribution of the two components is controlled by a specificity parameter{gamma} , and we show that the ability of a mutant to invade a resident population depends strongly on this parameter. When the carrying capacity is intrinsic, selection is almost exclusively for mutants with higher carrying capacity, while a shared carrying capacity yields selection purely on growth rate. This result has important implications for our understanding of niche construction, in particular the evolutionary dynamics of tumor growth.

Evolutionary Biology

Evidence for widespread positive and negative selection in coding and conserved noncoding regions of Capsella grandiflora

The extent that both positive and negative selection vary across different portions of plant genomes remains poorly understood. Here, we sequence whole genomes of 13 Capsella grandiflora individuals and quantify the amount of selection across the genome. Using an estimate of the distribution of fitness effects, we show that selection is strong in coding regions, but weak in most noncoding regions, with the exception of 5’ and 3’ untranslated regions (UTRs). However, estimates of selection in noncoding regions conserved across the Brassicaceae family show strong signals of selection. Additionally, we see reductions in neutral diversity around functional substitutions in both coding and conserved noncoding regions, indicating recent selective sweeps at these sites. Finally, using expression data from leaf tissue we show that genes that are more highly expressed experience stronger negative selection but comparable levels of positive selection to lowly expressed genes. Overall, we observe widespread positive and negative selection in coding and regulatory regions, but our results also suggest that both positive and negative selection in plant noncoding sequence are considerably rarer than in animal genomes.

Evolutionary Biology

Biochemical ‘Cambrian’ explosion-implosions: the generation and pruning of genetic codes

Tlusty's topological analysis of the genetic code suggests ecosystem changes in available metabolic free energy that predated the aerobic transition enabled a punctuated sequence of increasingly complex genetic codes and protein translators. These coevolved via a `Cambrian explosion' until, very early on, the ancestor of the present narrow spectrum of protein machineries became evolutionarily locked in at a modest level of fitness reflecting a modest embedding metabolic free energy ecology. Similar biochemical `Cambrian singularities' must have occurred at different scales and levels of organization on Earth, with competition or chance-selected outcomes frozen at a far earlier period than the physical bauplan Cambrian explosion. Other examples might include explosive variations in mechanisms of photosynthesis and subsequent oxygen metabolisms. Intermediate between Cambrian bauplan and genetic code, variants of both remain today, even after evolutionary pruning, often protected in specialized ecological niches. This suggests that, under less energetic astrobiological ecologies, a spectrum of less complicated reproductive codes may also survive in specialized niches.

Evolutionary Biology

The roles of standing genetic variation and evolutionary history in determining the evolvability of anti-predator strategies

Standing genetic variation and the historical environment in which that variation arises (evolutionary history) are both potentially significant determinants of a populations capacity for evolutionary response to a changing environment. We evaluated the relative importance of these two factors in influencing the evolutionary trajectories in the face of sudden environmental change. We used the open-ended digital evolution software Avida to examine how historic exposure to predation pressures, different levels of genetic variation, and combinations of the two, impact anti-predator strategies and competitive abilities evolved in the face of threats from new, invasive, predator populations. We show that while standing genetic variation plays some role in determining evolutionary responses, evolutionary history has the greater influence on a populations capacity to evolve effective anti-predator traits. This adaptability likely reflects the relative ease of repurposing existing, relevant genes and traits, and the broader potential value of the generation and maintenance of adaptively flexible traits in evolving populations.

Evolutionary Biology

Investigating speciation in face of polyploidization: what can we learn from approximate Bayesian computation approach?

Despite its importance in the diversification of many eucaryote clades, particularly plants, detailed genomic analysis of polyploid species is still in its infancy, with published analysis of only a handful of model species to date. Fundamental questions concerning the origin of polyploid lineages (e.g., auto- vs. allopolyploidy) and the extent to which polyploid genomes display disomic vs. polysomic vs. heterosomic inheritance are poorly resolved for most polyploids, not least because they have hitherto required detailed karyotypic analysis or the analysis of allele segregation at multiple loci in pedigrees or artificial crosses, which are often not practical for non-model species. However, the increasing availability of sequence data for non-model species now presents an opportunity to apply established approaches for the evolutionary analysis of genomic data to polyploid species complexes. Here, we ask whether approximate Bayesian computation (ABC), applied to sequence data produced by next-generation sequencing technologies from polyploid taxa, allows correct inference of the evolutionary and demographic history of polyploid lineages and their close relatives. We use simulations to investigate how the number of sampled individuals, the number of surveyed loci and their length affect the accuracy and precision of evolutionary and demographic inferences by ABC, including the mode of polyploidisation, mode of inheritance of polyploid taxa, the relative timing of genome duplication and speciation, and effective populations sizes of contributing lineages. We also apply the ABC framework we develop to sequence data from diploid and polyploidy species of the plant genus Capsella, for which we infer an allopolyploid origin for tetra C. bursa-pastoris {approx} 90,000 years ago. In general, our results indicate that ABC is a promising and powerful method for uncovering the origin and subsequent evolution of polyploid species.

Evolutionary Biology

Estimating the evolution of human life history traits in age-structured populations

I propose a method that estimates the selection response of all vital rates in an age-structured population. I assume that vital rates are determined by the additive genetic contributions of many loci. The method uses all relatedness information in the sample to inform its estimates of genetic parameters, via an MCMC Bayesian framework. One can use the results to estimate the selection response of any life history trait that is a function of the vital rates, including the age at first reproduction, total lifetime fertility, survival to adulthood, and others. This method closely ties the empirical analysis of life history evolution to dynamically complete models of natural selection, and therefore enjoys some theoretical advantages over other methods. I demonstrate the method on a simulated model of evolution with two age classes. Finally I discuss how the method can be extended to more complicated cases.

Evolutionary Biology

Differential gene expression and alternative splicing in insect immune specificity

Ecological studies routinely show genotype-genotype interactions between insects and their parasites. The mechanisms behind these interactions are not clearly understood. Using the bumblebee Bombus terrestris / trypanosome Crithidia bombi model system, we have carried out a transcriptome-wide analysis of gene expression and alternative splicing in bees during C. bombi infection. We have performed four analyses, 1) comparing gene expression in infected and non-infected bees 24 hours after infection by Crithidia bombi, 2) comparing expression at 24 and 48 hours after C.bombi infection, 3) searching for differential gene expression associated with the host-parasite genotype-genotype interaction at 24 hours after infection and 4) searching for alternative splicing associated with the host-parasite genotype-genotype interaction at 24 hours post infection. We found a large number of genes differentially regulated related to numerous canonical immune pathways. These genes include receptors, signaling pathways and effectors. We discovered a possible interaction between the peritrophic membrane and the insect immune system in defense against Crithidia. Most interestingly we found differential expression and alternative splicing of Dscam related transcripts and a novel immunoglobulin related gene Twitchin depends on the genotype-genotype interactions of the given bumblebee colony and Crithidia strain.

Evolutionary Biology

Migration and interaction in a contact zone: mtDNA variation among Bantu-speakers in southern Africa

Bantu speech communities expanded over large parts of sub-Saharan Africa within the last 4000-5000 years, reaching different parts of southern Africa 1200-2000 years ago. The Bantu languages subdivide in several major branches, with languages belonging to the Eastern and Western Bantu branches spreading over large parts of Central, Eastern, and Southern Africa. There is still debate whether this linguistic divide is correlated with a genetic distinction between Eastern and Western Bantu speakers. During their expansion, Bantu speakers would have come into contact with diverse local populations, such as the Khoisan hunter-gatherers and pastoralists of southern Africa, with whom they may have intermarried. In this study, we analyze complete mtDNA genome sequences from over 900 Bantu-speaking individuals from Angola, Zambia, Namibia, and Botswana to investigate the demographic processes at play during the last stages of the Bantu expansion. Our results show that most of these Bantu-speaking populations are genetically very homogenous, with no genetic division between speakers of Eastern and Western Bantu languages. Most of the mtDNA diversity in our dataset is due to different degrees of admixture with autochthonous populations. Only the pastoralist Himba and Herero stand out due to high frequencies of particular L3f and L3d lineages; the latter are also found in the neighboring Damara, who speak a Khoisan language and were foragers and small-stock herders. In contrast, the close cultural and linguistic relatives of the Herero and Himba, the Kuvale, are genetically similar to other Bantu-speakers. Nevertheless, as demonstrated by resampling tests, the genetic divergence of Herero, Himba, and Kuvale is compatible with a common shared ancestry with high levels of drift and differential female admixture with local pre-Bantu populations.

Evolutionary Biology

No evidence that natural selection has been less effective at removing deleterious mutations in Europeans than in West Africans

Non-African populations have experienced major bottlenecks in the time since their split from West Africans, which has led to the hypothesis that natural selection to remove weakly deleterious mutations may have been less effective in non-Africans. To directly test this hypothesis, we measure the per-genome accumulation of deleterious mutations across diverse humans. We fail to detect any significant differences, but find that archaic Denisovans accumulated non-synonymous mutations at a higher rate than modern humans, consistent with the longer separation time of modern and archaic humans. We also revisit the empirical patterns that have been interpreted as evidence for less effective removal of deleterious mutations in non-Africans than in West Africans, and show they are not driven by differences in selection after population separation, but by neutral evolution.

Evolutionary Biology

Mapping the structure of drosophilid behavior

Most animals possess the ability to actuate a vast diversity of movements, ostensibly constrained only by morphology and physics. In practice, however, a frequent assumption in behavioral science is that most of an animals activities can be described in terms of a small set of stereotyped motifs. Here we introduce a method for mapping the behavioral space of organisms, relying only upon the underlying structure of postural movement data to organize and classify behaviors. We find that six different drosophilid species each perform a mix of non-stereotyped actions and over one hundred hierarchically-organized, stereotyped behaviors. Moreover, we use this approach to compare these species behavioral spaces, systematically identifying subtle behavioral differences between closely-related species.

Evolutionary Biology

An experimentally determined evolutionary model dramatically improves phylogenetic fit

All modern approaches to molecular phylogenetics require a quantitative model for how genes evolve. Unfortunately, existing evolutionary models do not realistically represent the site-heterogeneous selection that governs actual sequence change. Attempts to remedy this problem have involved augmenting these models with a burgeoning number of free parameters. Here I demonstrate an alternative: experimental determination of a parameter-free evolutionary model via mutagenesis, functional selection, and deep sequencing. Using this strategy, I create an evolutionary model for influenza nucleoprotein that describes the gene phylogeny far better than existing models with dozens or even hundreds of free parameters. Emerging high-throughput experimental strategies such as the one employed here provide fundamentally new information that has the potential to transform the sensitivity of phylogenetic and genetic analyses.

Evolutionary Biology

Neanderthals had our de novo genes.

Gene duplication provides a profusion of raw material for evolutionary innovation (Lynch and Conery, 2000). While most duplicates rapidly become unrecognizable some, e.g., those that are immediately useful or those that after a period of relaxed selection gain unique roles, are retained and thereby expand a genomes protein-coding repertoire. Ohno (1970) famously remarked that without gene duplication, the creation of metazoans, vertebrates, and mammals from unicellular organisms would have been impossible. Such big leaps in evolution, he argued, required the creation of new gene loci with previously nonexistent functions (Taylor and Raes, 2004). Recently, another source of new genes has been recognized: Though rare, it seems clear that new genes can emerge from formerly non-coding DNA, the de novo protein coding genes (Zhao et al., 2014, and references therein).\n\nIn 2009 Knowles ...

Evolutionary Biology

Genetic drift opposes mutualism during spatial population expansion

Mutualistic interactions benefit both partners, promoting coexistence and genetic diversity. Spatial structure can promote cooperation, but spatial expansions may also make it hard for mutualistic partners to stay together, since genetic drift at the expansion front creates regions of low genetic and species diversity. To explore the antagonism between mutualism and genetic drift, we grew cross-feeding strains of the budding yeast S. cerevisiae on agar surfaces as a model for mutualists undergoing spatial expansions. By supplying varying amounts of the exchanged nutrients, we tuned strength and symmetry of the mutualistic interaction. Strong mutualism suppresses genetic demixing during spatial expansions and thereby maintains diversity, but weak or asymmetric mutualism is overwhelmed by genetic drift even when mutualism is still beneficial, slowing growth and reducing diversity. Theoretical modeling using experimentally measured parameters predicts the size of demixed regions and how strong mutualism must be to survive a spatial expansion.

Evolutionary Biology