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Hamilton's rule provides the cornerstone for our understanding of the evolution of all forms of social behavior, from altruism to spite, across all organisms, from viruses to humans. In contrast to the standard prediction from Hamilton's rule, recent studies have suggested that altruistic helping can be favored even if it does not benefit relatives, as long as it decreases the environmentally induced variance of their reproductive success ("altruistic bet-hedging"). However, previous predictions both rely on an approximation and focus on variance-reducing helping behaviors. We derived a version of Hamilton's rule that fully captures environmental variability. This shows that decreasing (or increasing) the variance in the absolute reproductive success of relatives does not have a consistent effect-it can either favor or disfavor the evolution of helping. We then empirically quantified the effect of helping on the variance in reproductive success across 15 species of cooperatively breeding birds. We found that a) helping did not consistently decrease the variance of reproductive success and often increased it, and b) the mean benefits of helping across environments consistently outweighed other variability components of reproductive success. Altogether, our theoretical and empirical results suggest that the effects of helping on the variability components of reproductive success have not played a consistent or strong role in favoring helping.

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Purpose: This study elucidates the relationship between similarity and ethnic psychological compatibility and its underlying psychological mechanisms. According to kin selection theory, similarity can promote ethnic psychological compatibility by enhancing psychological kinship and intergroup contact. Participants: and methods: A questionnaire survey was administered to 1523 participants from 25 ethnic groups in China. Data analysis was carried out via conditional process modelling. A multigroup comparison of mediation models between the ethnic majority and minorities was detected via the Stats Tools Package. Results: Our findings demonstrated that: (1) cultural and attitude similarity both showed a significant positive correlation with ethnic psychological compatibility; (2) psychological kinship and intergroup contact served as mediators in the relationship of attitude and cultural similarity with ethnic psychological compatibility and psychological kinship and intergroup contact were independent mediators, while psychological kinship-intergroup contact showed a significant serial mediation effect; (3) there were no significant differences in mediation effects between different ethnic groups. Conclusion: Our findings expand on kin selection theory and provide valuable paths for psychologically supporting ethnic psychological compatibility.

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Eusocial organisms typically live in colonies with one reproductive queen supported by thousands of sterile workers. It is widely believed that monogamous mating is a precondition for the evolution of eusociality. Here, we present a theoretical model that simulates a realistic scenario for the evolution of eusociality. In the model, mothers can evolve control over resource allocation to offspring, affecting offspring's body size. The offspring can evolve body-size-dependent dispersal, by which they disperse to breed or stay at the nest as helpers. We demonstrate that eusociality can evolve even if mothers are not strictly monogamous, provided that they can constrain their offspring's reproduction through manipulation. We also observe the evolution of social polymorphism with small individuals that help and larger individuals that disperse to breed. Our model unifies the traditional kin selection and maternal manipulation explanations for the evolution of eusociality and demonstrates that-contrary to current consensus belief-eusociality can evolve despite highly promiscuous mating.

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Male mice who are heterozygous for distorting and non-distorting alleles at the t-haplotype transmit the driving t-haplotype around 90% of the time - a drastic departure from Mendelian expectations. This selfish act comes at a cost. The mechanism underlying transmission distortion in this system causes severe sterility in males homozygous for the drive alleles, ultimately preventing its fixation. Curiously, many driving t-haplotypes also induce embryonic lethality in both sexes when homozygous; however, this is neither universal nor a necessity for this distortion mechanism. Charlesworth provided an adaptive explanation for the evolution of lethal t-haplotypes in a population segregating for distorting and non-distorting t alleles - if mothers compensate by replacing dead embryos with new offspring (or by transferring energy to surviving offspring), a recessive lethal can be favored because it effectively allows mothers the opportunity to trade in infertile males for potentially fertile offspring. This model, however, requires near complete reproductive compensation for the invasion of the lethal t-haplotype and produces an equilibrium frequency of lethal drivers well below what is observed in nature. We show that low levels of systemic inbreeding, which we model as brother-sister mating, allow lethal t-haplotypes to invade with much lower levels of reproductive compensation. Furthermore, inbreeding allows these lethal haplotypes to largely displace the ancestral male-sterile haplotypes. Our results show that together inbreeding and reproductive compensation move expected equilibria closer to observed haplotype frequencies in natural populations and occur under lower, potentially more reasonable, parameters.

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Behavioural and physiological resistance are key to slowing epidemic spread. We explore the evolutionary and epidemic consequences of their different costs for the evolution of tolerance that trades off with resistance. Behavioural resistance affects social cohesion, with associated group-level costs, while the cost of physiological resistance accrues only to the individual. Further, resistance, and the associated reduction in transmission, benefit susceptible hosts directly, whereas infected hosts only benefit indirectly, by reducing transmission to kin. We therefore model the coevolution of transmission-reducing resistance expressed in susceptible hosts with resistance expressed in infected hosts, as a function of kin association, and analyse the effect on population-level outcomes. Using parameter values for guppies, Poecilia reticulata, and their gyrodactylid parasites, we find that: (1) either susceptible or infected hosts should invest heavily in resistance, but not both; (2) kin association drives investment in physiological resistance more strongly than in behavioural resistance; and (3) even weak levels of kin association can favour altruistic infected hosts that invest heavily in resistance (versus selfish tolerance), eliminating parasites. Overall, our finding that weak kin association affects the coevolution of infected and susceptible investment in both behavioural and physiological resistance suggests that kin selection may affect disease dynamics across systems.

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The growth and success of many bacteria appear to rely on a stunning range of cooperative behaviours. But what is cooperation and how is it studied?

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Upon uptake of toxins, insects launch a detoxification program. This program is deployed in multiple organs and cells to raise their tolerance against the toxin. The molecular mechanisms of this program inside the insect body have been studied and understood in detail. Here, we report on a yet unexplored extra-corporeal detoxification of insecticides in Drosophila melanogaster. Wild-type D. melanogaster incubated with DDT, a contact insecticide, in a closed environment died as expected. However, incubation of a second cohort in the same environment after removal of the dead flies was not lethal. The effect was significantly lower if the flies of the two cohorts were unrelated. Incubation assays with Chlorpyrifos, another contact insecticide, yielded identical results, while incubation assays with Chlorantraniliprole, again a contact insecticide, was toxic for the second cohort of flies. A cohort of flies incubated in a DDT environment after an initial incubation of a honeybee survived treatment. Together, our data suggest that insects including Apis mellifera and D. melanogaster have the capacity to modify their proximate environment. Consequently, in their ecological niche, following individuals might be saved from intoxication thereby facilitating colonisation of an attractive site.

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A long-standing problem in evolutionary theory is to clarify in what sense (if any) natural selection cumulatively improves the design of organisms. Various concepts, such as fitness and inclusive fitness, have been proposed to resolve this problem. In addition, there have been attempts to replace the original problem with more tractable questions, such as whether a given gene or trait is favored by selection. Here, we ask what theoretical properties the concept fitness should possess to encapsulate the improvement criterion required to talk meaningfully about adaptive evolution. We argue that natural selection tends to shape phenotypes based on the causal properties of individuals and that this tendency is, therefore, best captured by a fitness concept that focuses on these properties. We highlight a fitness concept that meets this role under broad conditions but requires adjustments in our conceptual understanding of adaptive evolution. These adjustments combine elements of Dawkinsian gene selectionism and Egbert Leigh's "parliament of genes."

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Greenbeards are selfish genetic elements that make their bearers behave either altruistically towards individuals bearing similar greenbeard copies or harmfully towards individuals bearing different copies. They were first proposed by W. D. Hamilton over 50 yr ago, to illustrate that kin selection may operate at the level of single genes. Examples of greenbeards have now been reported in a wide range of taxa, but they remain undocumented in plants. In this paper, we discuss the theoretical likelihood of greenbeard existence in plants. We then question why the greenbeard concept has never been applied to plants and speculate on how hypothetical greenbeards could affect plant-plant interactions. Finally, we point to different research directions to improve our knowledge of greenbeards in plants.

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Much research on the evolution of altruism via kin selection, group selection, and reciprocity focuses on the role of a single locus or quantitative trait. Very few studies have explored how linked selection, or selection at loci neighboring an altruism locus, impacts the evolution of altruism. While linked selection can decrease the efficacy of selection at neighboring loci, it might have other effects including promoting selection for altruism by increasing relatedness in regions of low recombination. Here, we used population genetic simulations to study how negative selection at linked loci, or background selection, affects the evolution of altruism. When altruism occurs between full siblings, we found that background selection interfered with selection on the altruistic allele, increasing its fixation probability when the altruistic allele was disfavored and reducing its fixation when the allele was favored. In other words, background selection has the same effect on altruistic genes in family-structured populations as it does on other, nonsocial, genes. This contrasts with prior research showing that linked selective sweeps can favor the evolution of cooperation, and we discuss possibilities for resolving these contrasting results.

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Individuals delay natal dispersal for many reasons. There may be no place to disperse to; immediate dispersal or reproduction may be too costly; immediate dispersal may mean that the individual and their relatives miss the benefits of group living. Understanding the factors that lead to the evolution of delayed dispersal is important because delayed dispersal sets the stage for complex social groups and social behavior. Here, we study the evolution of delayed dispersal when the quality of the local environment is improved by greater numbers of individuals (e.g., safety in numbers). We assume that individuals who delay natal dispersal also expect to delay personal reproduction. In addition, we assume that improved environmental quality benefits manifest as changes to fecundity and survival. We are interested in how do the changes in these life-history features affect delayed dispersal. We use a model that ties evolution to population dynamics. We also aim to understand the relationship between levels of delayed dispersal and the probability of establishing as an independent breeder (a population-level feature) in response to changes in life-history details. Our model emphasizes kin selection and considers a sexual organism, which allows us to study parent-offspring conflict over delayed dispersal. At evolutionary equilibrium, fecundity and survival benefits of group size or quality promote higher levels of delayed dispersal over a larger set of life histories with one exception. The exception is for benefits of increased group size or quality reaped by the individuals who delay dispersal. There, the increased benefit does not change the life histories supporting delay dispersal. Next, in contrast to previous predictions, we find that a low probability of establishing in a new location is not always associated with a higher incidence of delayed dispersal. Finally, we find that increased personal benefits of delayed dispersal exacerbate the conflict between parents and their offspring. We discuss our findings in relation to previous theoretical and empirical work, especially work related to cooperative breeding.

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According to Michener's paradox, most altruistic groups in nature should be small and large groups should not exist. This is because per capita productivity is thought to decrease as groups get larger, meaning that the share of indirect fitness available to each group member declines, which favours dispersal. The empirical evidence for a decrease in per capita productivity is contradictory, however, and limited to the social Hymenoptera. I report that per capita reproductive success decreased with increasing group size across 26 cooperatively breeding bird species. Small groups comprising two or three individuals were the most common (79% of 16,101 groups), and these had the highest per capita reproductive success. This close fit between per capita reproductive success and the distribution of group sizes in nature suggests that it may indeed be difficult for large groups to evolve through indirect fitness benefits alone.

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AbstractIn cooperative breeding systems, inclusive fitness theory predicts that nonbreeding helpers more closely related to the breeders should be more willing to provide costly alloparental care and thus have more impact on breeder fitness. In the red-cockaded woodpecker (Dryobates borealis), most helpers are the breeders' earlier offspring, but helpers do vary within groups in both relatedness to the breeders (some even being unrelated) and sex, and it can be difficult to parse their separate impacts on breeder fitness. Moreover, most support for inclusive fitness theory has been positive associations between relatedness and behavior rather than actual fitness consequences. We used functional linear models to evaluate the per capita effects of helpers of different relatedness on eight breeder fitness components measured for up to 41 years at three sites. In support of inclusive fitness theory, helpers more related to the breeding pair made greater contributions to six fitness components. However, male helpers made equal contributions to increasing prefledging survival regardless of relatedness. These findings suggest that both inclusive fitness benefits and other direct benefits may underlie helping behaviors in the red-cockaded woodpecker. Our results also demonstrate the application of an underused statistical approach to disentangle a complex ecological phenomenon.

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Because plumage coloration is frequently involved in sexual selection, for both male and female mate choice, birds with aberrant plumage should have fewer mating opportunities and thus lower reproductive output. Here we report an Eastern Bluebird (Sialia sialis) female with a brown phenotype that raised a brood of four chicks to fledging. The brown female and her mate were only related to their social offspring to the second degree and one of the offspring was a half-sibling. We propose four family tree scenarios and discuss their implications (e.g., extra-pair paternity, conspecific brood parasitism). Regardless of the tree, the brown female was able to find a mate, which may have been facilitated by the bottleneck created by the severe snowstorms in February 2021.

Parce que la coloration du plumage est fréquemment impliquée dans la sélection sexuelle, tant pour le choix du partenaire mâle que femelle, les oiseaux avec un plumage aberrant devraient avoir moins d'opportunités d'accouplement et donc un rendement de reproduction réduit. Nous rapportons ici le cas d'une femelle Merlebleu de l'Est (Sialia sialis) avec un phénotype brun qui a élevé une nichée de quatre poussins jusqu'à l'envol. La femelle brune et son partenaire n'étaient liés à leur progéniture sociale qu'au second degré et l'un des poussins était un demi­frère ou une demi­sœur. Nous proposons quatre scenarios d'arbre généalogique et discutons de leurs implications (par exemple, paternité hors couple, parasitisme conspécifique), mais quel que soit l'arbre, la femelle brune a pu trouver un partenaire, ce qui a peut­être été facilité par le goulot d'étranglement créé par les fortes tempêtes de neige en février 2021.

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Fisher's geometric model provides a powerful tool for making predictions about key properties of Darwinian adaptation. Here, I apply the geometric model to predict differences between the evolution of altruistic versus nonsocial phenotypes. I recover Kimura's prediction that probability of fixation is greater for mutations of intermediate size, but I find that the effect size that maximises probability of fixation is relatively small in the context of altruism and relatively large in the context of nonsocial phenotypes, and that the overall probability of fixation is lower for altruism and is higher for nonsocial phenotypes. Accordingly, the first selective substitution is expected to be smaller, and to take longer, in the context of the evolution of altruism. These results strengthen the justification for employing streamlined social evolutionary methodologies that assume adaptations are underpinned by many genes of small effect.

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Bacteria cooperate by working collaboratively to defend their colonies, share nutrients, and resist antibiotics. Nevertheless, our understanding of these remarkable behaviours primarily comes from studying a few well-characterized species. Consequently, there is a significant gap in our understanding of microbial social traits, particularly in natural environments. To address this gap, we can use bioinformatic tools to identify genes that control cooperative or otherwise social traits. Existing tools address this challenge through two approaches. One approach is to identify genes that encode extracellular proteins, which can provide benefits to neighbouring cells. An alternative approach is to predict gene function using annotation tools. However, these tools have several limitations. Not all extracellular proteins are cooperative, and not all cooperative behaviours are controlled by extracellular proteins. Furthermore, existing functional annotation methods frequently miss known cooperative genes. We introduce SOCfinder as a new tool to find bacterial genes that control cooperative or otherwise social traits. SOCfinder combines information from several methods, considering if a gene is likely to [1] code for an extracellular protein [2], have a cooperative functional annotation, or [3] be part of the biosynthesis of a cooperative secondary metabolite. We use data on two extensively-studied species (P. aeruginosa and B. subtilis) to show that SOCfinder is better at finding known cooperative genes than existing tools. We also use theory from population genetics to identify a signature of kin selection in SOCfinder cooperative genes, which is lacking in genes identified by existing tools. SOCfinder opens up a number of exciting directions for future research, and is available to download from https://github.com/lauriebelch/SOCfinder.

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The evolution of cooperation depends on two crucial overarching factors: relatedness, which describes the extent to which the recipient shares genes in common with the actor; and quality, which describes the recipient's basic capacity to transmit genes into the future. While most research has focused on relatedness, there is a growing interest in understanding how quality modulates the evolution of cooperation. However, the impact of inheritance of quality on the evolution of cooperation remains largely unexplored, especially in spatially structured populations. Here, we develop a mathematical model to understand how inheritance of quality, in the form of social status, influences the evolution of helping and harming within social groups in a viscous-population setting. We find that: (1) status-reversal transmission, whereby parental and offspring status are negatively correlated, strongly inhibits the evolution of cooperation, with low-status individuals investing less in cooperation and high-status individuals being more prone to harm; (2) transmission of high status promotes offspring philopatry, with more cooperation being directed towards the higher-dispersal social class; and (3) fertility inequality and inter-generational status inheritance reduce within-group conflict. Overall, our study highlights the importance of considering different mechanisms of phenotypic inheritance, including social support, and their potential interactions in shaping animal societies.

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Social behaviours are typically modelled using neighbour-modulated fitness, which focuses on individuals having their fitness altered by neighbours. However, these models are either interpreted using inclusive fitness, which focuses on individuals altering the fitness of neighbours, or not interpreted at all. This disconnect leads to interpretational mistakes and obscures the adaptive significance of behaviour. We bridge this gap by presenting a systematic methodology for constructing inclusive-fitness models. We find a behaviour's 'inclusive-fitness effect' by summing primary and secondary deviations in reproductive value. Primary deviations are the immediate result of a social interaction; for example, the cost and benefit of an altruistic act. Secondary deviations are compensatory effects that arise because the total reproductive value of the population is fixed; for example, the increased competition that follows an altruistic act. Compared to neighbour-modulated fitness methodologies, our approach is often simpler and reveals the model's inclusive-fitness narrative clearly. We implement our methodology first in a homogeneous population, with supplementary examples of help under synergy, help in a viscous population and Creel's paradox. We then implement our methodology in a class-structured population, where the advantages of our approach are most evident, with supplementary examples of altruism between age classes, and sex-ratio evolution.

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BACKGROUND: Conventional wisdom in evolutionary theory considers aging as a non-selected byproduct of natural selection. Based on this, conviction aging was regarded as an inevitable phenomenon. It was also thought that in the wild organisms tend to die from diseases, predation and other accidents before they could reach the time when senescence takes its course. Evidence has accumulated, however, that aging is not inevitable and there are organisms that show negative aging even. Furthermore, old age does play a role in the deaths of many different organisms in the wild also. The hypothesis of programmed aging posits that a limited lifespan can evolve as an adaptation (i.e., positively selected for) in its own right, partly because it can enhance evolvability by eliminating "outdated" genotypes. A major shortcoming of this idea is that non-aging sexual individuals that fail to pay the demographic cost of aging would be able to steal good genes by recombination from aging ones. RESULTS: Here, we show by a spatially explicit, individual-based simulation model that aging can positively be selected for if a sufficient degree of kin selection complements directional selection. Under such conditions, senescence enhances evolvability because the rate of aging and the rate of recombination play complementary roles. The selected aging rate is highest at zero recombination (clonal reproduction). In our model, increasing extrinsic mortality favors evolved aging by making up free space, thereby decreasing competition and increasing drift, even when selection is stabilizing and the level of aging is set by mutation-selection balance. Importantly, higher extrinsic mortality is not a substitute for evolved aging under directional selection either. Reduction of relatedness decreases the evolved level of aging; chance relatedness favors non-aging genotypes. The applicability of our results depends on empirical values of directional and kin selection in the wild. CONCLUSIONS: We found that aging can positively be selected for in a spatially explicit population model when sufficiently strong directional and kin selection prevail, even if reproduction is sexual. The view that there is a conceptual link between giving up clonal reproduction and evolving an aging genotype is supported by computational results.

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