Kin Selection Research Articles

Calculating Relatedness: A Pedigree of Definitions.

Biology can be viewed from both an organismal and a genic perspective. A good example is W.D. Hamilton's work on inclusive fitness and kin selection, which puts relatedness at the heart of our understanding of social behavior. Relatedness mediates how much an actor should value a specific behavior's effect on a relative compared to the cost incurred to itself. Despite its key explanatory role, relatedness is also a concept marred with misunderstanding. Part of the problem has been that the term has been used in different ways by different people. To help address this, we survey the history of how relatedness has been formally modeled, paying particular attention to how it is conceptualized from both a gene-centric and an organism-centric point of view.

Cooperative and Solitary Mobbing to Defend Offspring in Wild Gray Mouse Lemurs, Microcebus murinus

ABSTRACTGray mouse lemur, Microcebus murinus, mothers either cooperatively breed plurally with kin or breed solitarily. We describe the first observations of cooperative and solitary mobbing to defend offspring by wild cooperative breeding gray mouse lemur mothers in Kirindy Forest, Western Madagascar. We observed four groups of cooperatively breeding mothers and their offspring daily between 18:00 and 04:00 from January 1 to 15, 2023. Cooperative mobbing was observed twice, and solitary mobbing by a single cooperative breeding mother was observed once. There was one Malagasy tree boa, Sanzinia madagascariensis, per mobbing event. Mothers solitarily mobbed by directly approaching within 1 m of the boa, walking and changing distance while within 1 m of the boa. In addition to the solitary mobbing behaviors, cooperatively mobbing mothers approached the boa together from the same or different directions or alternated, and they gathered around the boa. Mothers collaborated in the context of offspring defense from predators and performed different complementary tasks: mobbing while another provided alloparental care by guarding the offspring of the association, that is, babysitting. Mothers performed similar mobbing behaviors and occasionally synchronized in time or coordinate in time and in space to approach from different directions and gather around the boa. Mothers did not come in contact with the boas during mobbing but kept a distance of at least 20 cm. On two mobbing events, the boa rose up with its head facing the mobbing mothers and climbed down the tree after the mobbing. Cooperative mobbing to defend offspring likely evolved by kin selection but reciprocity and an interaction between kin selection and reciprocity cannot be ruled out, since gray mouse lemur mothers cooperatively breed with kin and basically allonurse reciprocally.

Disentangling the Factors Selecting for Unicellular Programmed Cell Death.

AbstractThe widespread occurrence of genetically programmed cell death (PCD) in unicellular species poses an evolutionary puzzle. While kin selection theory predicts that the fitness benefits of cell suicide must be preferentially directed toward genetic relatives, it does not predict the nature of these benefits. Furthermore, cell suicide must be conditionally expressed, leaving open the question of what conditions optimally regulate expression. Here we formalize several verbal hypotheses for the ecological function of unicellular PCD. We show that self-sacrifice by healthy cells cannot evolve. Instead, PCD evolution requires that damaged cells sense impending death and then (1) expedite this death to spare resources for groupmates, (2) prepare cellular contents so that necrotic toxins are not released upon death, or initiate autolysis in order to (3) release beneficial compounds or (4) release anticompetitior toxins. The prerequisite ability to predict death is a severe cell biological constraint as well as an ecological constraint that restricts PCD evolution to species with specific sources of mortality. We show that the specific type of PCD that will evolve, though, differs on the basis of a species' ecology, life history, and genetic structure.

Altruism and natural selection in a variable environment

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.

Maternal manipulation of offspring size can trigger the evolution of eusociality in promiscuous species

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.

Similarity enhances psychological compatibility: Serial mediation effect of psychological kinship and intergroup contact

PurposeThis 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. Participantsand 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. ResultsOur 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. ConclusionOur findings expand on kin selection theory and provide valuable paths for psychologically supporting ethnic psychological compatibility.

A Theoretical Comparison of Alternative Male Mating Strategies in Cephalopods and Fishes.

We used computer simulations of growth, mating and death of cephalopods and fishes to explore the effect of different life-history strategies on the relative prevalence of alternative male mating strategies. Specifically, we investigated the consequences of single or multiple matings per lifetime, mating strategy switching, cannibalism, resource stochasticity, and altruism towards relatives. We found that a combination of single (semelparous) matings, cannibalism and an absence of mating strategy changes in one lifetime led to a more strictly partitioned parameter space, with a reduced region where the two mating strategies co-exist in similar numbers. Explicitly including Hamilton's rule in simulations of the social system of a Cichlid led to an increase of dominant males, at the expense of both sneakers and dwarf males ("super-sneakers"). Our predictions provide general bounds on the viable ratios of alternative male mating strategies with different life-histories, and under possibly rapidly changing ecological situations.

Kin Altruism Questionnaire – theoretical foundations, construction of the tool and its psychometric characteristics

Kin Altruism Questionnaire – theoretical foundations, construction of the tool and its psychometric characteristics

Evaluation of breeding strategies to reduce the inbreeding rate in the Friesian horse population: Looking back and moving forward.

In the past, small population sizes and unequal ancestor contributions have resulted in high inbreeding rates (ΔF) in the Friesian horse. Two decades ago, the studbook implemented a mating quota and started publishing individual kinships and reduced ΔF below 1% per generation. However, since then, the breeding population size has decreased and this raises the question whether current breeding strategies are sufficient to keep ΔF below desired rates. The aim of this study was to (1) reflect on past inbreeding trends and their main determinants, using pedigree analysis and (2) evaluate the effectiveness of the current and additional breeding strategies using stochastic simulations. We estimated the current ΔF (2013-2022) at 0.72% per generation. While the total contribution of the top 10 sires to the number of offspring per year has decreased from 75% in 1980 to 35% in 2022, this was mainly due to an increased number of approved studbook sires, and not due to more equalized contributions among sires. Of the simulated breeding strategies, selecting only breeding stallions with a below average mean kinship (i.e., "mean kinship selection") was most effective to decrease ΔF (from 0.66% to 0.33%). Increasing the number of breeding sires only had an effect when also a mating quota was applied. However, its effect remained limited. For example, a ~1.5 fold increase, combined with a mating quota of 80 offspring per sire per year, reduced ΔF from 0.55% to 0.51%. When increasing the number of breeding mares, a practically unfeasible large increase was needed for a meaningful reduction in ΔF (e.g. twice as many mares were needed to reduce ΔF from 0.66% to 0.56%). Stratified mating quotas, a novel approach in which we assigned each sire a mating quota (of 60, 80, 100 or 120 offspring per year) based on its mean kinship to recently born foals, resulted in a lower ΔF (0.43%) than a general mating quota of 90 offspring per sire per year (0.55%). Overall, while the current ΔF is below 1%, we recommend to implement additional strategies to further reduce ΔF below 0.5% in the Friesian horse population. For this breed and similar populations, we recommend to focus on breeding strategies based on kinship levels to effectively reduce ΔF.

Group-selection via aggregative propagule-formation enables cooperative multicellularity in an individual based, spatial model.

The emergence of multicellularity is one of the major transitions in evolution that happened multiple times independently. During aggregative multicellularity, genetically potentially unrelated lineages cooperate to form transient multicellular groups. Unlike clonal multicellularity, aggregative multicellular organisms do not rely on kin selection instead other mechanisms maintain cooperation against cheater phenotypes that benefit from cooperators but do not contribute to groups. Spatiality with limited diffusion can facilitate group selection, as interactions among individuals are restricted to local neighbourhoods only. Selection for larger size (e.g. avoiding predation) may facilitate the emergence of aggregation, though it is unknown, whether and how much role such selection played during the evolution of aggregative multicellularity. We have investigated the effect of spatiality and the necessity of predation on the stability of aggregative multicellularity via individual-based modelling on the ecological timescale. We have examined whether aggregation facilitates the survival of cooperators in a temporally heterogeneous environment against cheaters, where only a subset of the population is allowed to periodically colonize a new, resource-rich habitat. Cooperators constitutively produce adhesive molecules to promote aggregation and propagule-formation while cheaters spare this expense to grow faster but cannot aggregate on their own, hence depending on cooperators for long-term survival. We have compared different population-level reproduction modes with and without individual selection (predation) to evaluate the different hypotheses. In a temporally homogeneous environment without propagule-based colonization, cheaters always win. Predation can benefit cooperators, but it is not enough to maintain the necessary cooperator amount in successive dispersals, either randomly or by fragmentation. Aggregation-based propagation however can ensure the adequate ratio of cooperators-to-cheaters in the propagule and is sufficient to do so even without predation. Spatiality combined with temporal heterogeneity helps cooperators via group selection, thus facilitating aggregative multicellularity. External stress selecting for larger size (e.g. predation) may facilitate aggregation, however, according to our results, it is neither necessary nor sufficient for aggregative multicellularity to be maintained when there is effective group-selection.

Juvenile mortality and sibling replacement: a kin selection approach

Abstract Mortality generally is higher around birth and then progressively declines through the juvenile stage. In species where offspring depend upon their parents during maturation, a factor behind this mortality decline could be sibling replacement: offspring sacrifice their survival to benefit future or present siblings as early as possible in order to minimize losses in parental investment. Here, we propose a kin-selection model of sibling replacement. Theoretical analysis of the model and its application to demographic data of mammals suggest that sibling replacement consistently generates a selective incentive for increasing juvenile mortality at early ages when this mortality increment is the result of positive selection for juvenile altruism within the nuclear family. The model highlights how sibling replacement goes beyond optimal allocation of parental resources into dependents and can provoke greater mortality closer to birth also in response to a more favorable ratio of actors to recipients of altruism among siblings.

Associating with kin selects for disease resistance and against tolerance.

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.

Reproductive Cessation and Post-Reproductive Lifespan in Honeybee Workers.

The post-reproductive lifespan is an evolutionary enigma because the cessation of reproduction in animals seems contrary to the maximization of Darwinian fitness. Several theories aim to explain the evolution of menopause, one of which suggests that females of a certain age receive more fitness benefits via indirect selection (kin selection) than they would directly from continuing reproduction. Post-reproductive lifespans are not very common in nature but have been described in humans, nonhuman primates, a few species of toothed whales, guppies, and in some insect societies consisting of clonal colony members, such as aphid and ant societies. Here, we provide evidence that menopause also exists in honeybee societies. Our study shows that workers with a short life expectancy (older and/or injured workers) invest fewer resources and less time in their own reproduction than workers with a long life expectancy (younger and/or uninjured workers), even if their colony is hopelessly queenless. These results are consistent with the kin selection explanation for the evolution of menopause and help us understand the net effects of relatedness and social cooperation in animals.

Proximity and preening in captive Humboldt penguins

Group-living animals, including penguins, exhibit affiliative behaviors such as grooming (preening) and proximity. Such behaviors in non-primate animals have been less studied than those in primates. Our research focused on 20 identifiable Humboldt penguins in a zoo, analyzing kin relationships and reciprocity in preening and proximity by employing a 5-minute scan sampling method to observe and record individual behavior. Our findings revealed that preening and proximity were more prevalent among mate pairs. However, among non-mate pairs, such behaviors were more commonly observed between siblings and parent-offspring pairs. Notably, the individuals preened on each other simultaneously in all instances. This study highlights the potential influence of kin selection in shaping the affiliative behavior of penguins. Additionally, our findings indicate that penguins gain benefits from mutual preening. This study contributes to our understanding of social behaviors in non-primate species and emphasizes the need for further comparative studies of various animal taxa to elucidate the evolution of sociality.

Inferring stochastic group interactions within structured populations via coupled autoregression

The internal behaviour of a population is an important feature to take account of when modelling its dynamics. In line with kin selection theory, many social species tend to cluster into distinct groups in order to enhance their overall population fitness. Temporal interactions between populations are often modelled using classical mathematical models, but these sometimes fail to delve deeper into the, often uncertain, relationships within populations. Here, we introduce a stochastic framework that aims to capture the interactions of animal groups and an auxiliary population over time. We demonstrate the model’s capabilities, from a Bayesian perspective, through simulation studies and by fitting it to predator–prey count time series data. We then derive an approximation to the group correlation structure within such a population, while also taking account of the effect of the auxiliary population. We finally discuss how this approximation can lead to ecologically realistic interpretations in a predator–prey context. This approximation also serves as verification to whether the population in question satisfies our various assumptions. Our modelling approach will be useful for empiricists for monitoring groups within a conservation framework and also theoreticians wanting to quantify interactions, to study cooperation and other phenomena within social populations.

The Exploring Mechanisms of the Formation of Altruistic Behaviors

Altruism is essential to the world. If people do something on account of caring for others more, it is altruistic. Evolution motivates kin altruism. Reciprocal altruism is essential to human society. There also exists altruism that has nothing to do with gene correlation, like vernacular altruism. Different kinds of altruism have different motivations. The glow effect exists because people want to experience again the warm glow they feel when being altruistic, though the neural base needs to be studied. A costly long-term investment is a more complex altruistic behavior in the real world. It can be explained by selective investment theory, which discusses the importance of relationships. What’s more, altruism has its neurological base. The amygdala volume of people with altruistic trait may be more significant. They may also have more excellent neural activity in regions concerning caring and empathy, like midbrain periaqueductal gray and anterior insular. These neurological findings also lay the foundation for some people to be more altruistic than others. Future research can enlarge the investigation on the practical use of altruism, like discovering how to activate people’s altruistic behaviors and what people can do to cultivate and preserve altruism.

Genes for cooperation are not more likely to be carried by plasmids.

Cooperation is prevalent across bacteria, but risks being exploited by non-cooperative cheats. Horizontal gene transfer, particularly via plasmids, has been suggested as a mechanism to stabilize cooperation. A key prediction of this hypothesis is that genes which are more likely to be transferred, such as those on plasmids, should be more likely to code for cooperative traits. Testing this prediction requires identifying all genes for cooperation in bacterial genomes. However, previous studies used a method which likely misses some of these genes for cooperation. To solve this, we used a new genomics tool, SOCfinder, which uses three distinct modules to identify all kinds of genes for cooperation. We compared where these genes were located across 4648 genomes from 146 bacterial species. In contrast to the prediction of the hypothesis, we found no evidence that plasmid genes are more likely to code for cooperative traits. Instead, we found the opposite-that genes for cooperation were more likely to be carried on chromosomes. Overall, the vast majority of genes for cooperation are not located on plasmids, suggesting that the more general mechanism of kin selection is sufficient to explain the prevalence of cooperation across bacteria.

Evolution of delayed dispersal with group size effect and population dynamics

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.

Siderophores promote cooperative interspecies and intraspecies cross-protection against antibiotics in vitro.

The antibiotic cefiderocol hijacks iron transporters to facilitate its uptake and resists β-lactamase degradation. While effective, resistance has been detected clinically with unknown mechanisms. Here, using experimental evolution, we identified cefiderocol resistance mutations in Pseudomonas aeruginosa. Resistance was multifactorial in host-mimicking growth media, led to multidrug resistance and paid fitness costs in cefiderocol-free environments. However, kin selection drove some resistant populations to cross-protect susceptible individuals from killing by increasing pyoverdine secretion via a two-component sensor mutation. While pyochelin sensitized P. aeruginosa to cefiderocol killing, pyoverdine and the enterobacteria siderophore enterobactin displaced iron from cefiderocol, preventing uptake by susceptible cells. Among 113 P. aeruginosa intensive care unit clinical isolates, pyoverdine production directly correlated with cefiderocol tolerance, and high pyoverdine producing isolates cross-protected susceptible P. aeruginosa and other Gram-negative bacteria. These in vitro data show that antibiotic cross-protection can occur via degradation-independent mechanisms and siderophores can serve unexpected protective cooperative roles in polymicrobial communities.

Greenbeards in plants?

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.