Kin competition,natal dispersal and the moulding of senescence by natural selection

Most theoretical models for the evolution of senescence have assumed a very large, well mixed population. Here, we investigate how limited dispersal and kin competition might influence the evolution of ageing by deriving indicators of the force of selection, similar to Hamilton (Hamilton 1966 J. Theor. Biol. 12, 12–45). Our analytical model describes how the strength of selection on survival and fecundity changes with age in a patchy population, where adults are territorial and a fraction of juveniles disperse between territories. Both parent–offspring competition and sib competition then affect selection on age-specific life-history traits. Kin competition reduces the strength of selection on survival. Mutations increasing mortality in some age classes can even be favoured by selection, but only when fecundity deteriorates rapidly with age. Population structure arising from limited dispersal however selects for a broader distribution of reproduction over the lifetime, potentially slowing down reproductive senescence. The antagonistic effects of limited dispersal on age schedules of fecundity and mortality cast doubts on the generality of conditions allowing the evolution of ‘suicide genes’ that increase mortality rates without other direct pleiotropic effects. More generally, our model illustrates how limited dispersal and social interactions can indirectly produce patterns of antagonistic pleiotropy affecting vital rates at different ages.     

Kin selection and the evolution of sexual conflict

Males and females do not always share the same evolutionary interests. This is particularly true in the case of multiple mating, where male–male competition can often lead to adaptations that are harmful to the female, and females can evolve counter adaptations to reduce the benefits males gain from such traits. Although social evolution has made substantial progress from kin selection theory, most studies of sexual conflict have ignored the effects of genetic relatedness. Here, I use a model of male harm and female resistance to investigate how kin selection affects the evolution of sexual conflict. Building on models of social evolution, I show that relatedness inhibits sexual conflict, in terms of male harm, whereas it has no effect on the evolution female resistance. This study examines a previously neglected mechanism that can potentially help to resolve sexual conflict over mating and highlights the potential importance of considering relatedness in empirical studies of sexual conflict.     

Asymmetry within social groups: division of labour and intergroup competition

Social animals vary in their ability to compete with group members over shared resources and also vary in their cooperative efforts to produce these resources. Competition among groups can promote within‐group cooperation, but many existing models of intergroup cooperation do not explicitly account for observations that group members invest differentially in cooperation and that there are often within‐group competitive or power asymmetries. We present a game theoretic model of intergroup competition that investigates how such asymmetries affect within‐group cooperation. In this model, group members adopt one of two roles, with relative competitive efficiency and the number of individuals varying between roles. Players in each role make simultaneous, coevolving decisions. The model predicts that although intergroup competition increases cooperative contributions to group resources by both roles, contributions are predominantly from individuals in the less competitively efficient role, whereas individuals in the more competitively efficient role generally gain the larger share of these resources. When asymmetry in relative competitive efficiency is greater, a group's per capita cooperation (averaged across both roles) is higher, due to increased cooperation from the competitively inferior individuals. For extreme asymmetry in relative competitive efficiency, per capita cooperation is highest in groups with a single competitively superior individual and many competitively inferior individuals, because the latter acquiesce and invest in cooperation rather than within‐group competition. These predictions are consistent with observed features of many societies, such as monogynous Hymenoptera with many workers and caste dimorphism.     

The robustness of the weak selection approximation for the evolution of altruism against strong selection

The weak selection approximation of population genetics has made possible the analysis of social evolution under a considerable variety of biological scenarios. Despite its extensive usage, the accuracy of weak selection in predicting the emergence of altruism under limited dispersal when selection intensity increases remains unclear. Here, we derive the condition for the spread of an altruistic mutant in the infinite island model of dispersal under a Moran reproductive process and arbitrary strength of selection. The simplicity of the model allows us to compare weak and strong selection regimes analytically. Our results demonstrate that the weak selection approximation is robust to moderate increases in selection intensity and therefore provides a good approximation to understand the invasion of altruism in spatially structured population. In particular, we find that the weak selection approximation is excellent even if selection is very strong, when either migration is much stronger than selection or when patches are large. Importantly, we emphasize that the weak selection approximation provides the ideal condition for the invasion of altruism, and increasing selection intensity will impede the emergence of altruism. We discuss that this should also hold for more complicated life cycles and for culturally transmitted altruism. Using the weak selection approximation is therefore unlikely to miss out on any demographic scenario that lead to the evolution of altruism under limited dispersal.     

Bidirectional transport of organelles: unity and struggle of opposing motors

Bidirectional transport along microtubules is ensured by opposing motor proteins: cytoplasmic dynein that drives cargo to the minus-ends and various kinesins that generally move to the plus-ends of microtubules. Regulation of motor proteins that are simultaneously bound to the same organelle is required to maintain directional transport and prevent pausing of cargo pulled away by motors of opposite polarity. Debates of the recent decade have been focused on two possible mechanisms of such regulation: (i) coordination, which implies that only one type of motors is active at a given time, and (ii) tug-of-war, which assumes that both motors are active at the same time and that direction of transport depends on the outcome of motor's confrontation. The initial idea of coordination has been challenged by observations of simultaneous activity of plus- and minus-end-directed motors applied to the same cargo. Analysis of the available data indicates that coordination and tug-of-war theories rather complement than contradict each other: cargo interacts with two teams of active motors, the resulting direction and the winner team are determined by coordination complexes, but the activity of the loser team is never completely inhibited and remains at some background level. Such persisting activity might enhance the overall efficiency of transport by increasing processivity or helping to overcome the obstacles on microtubule track.     

Kin selection,species richness and community

Can evolutionary and ecological dynamics operating at one level of the biological hierarchy affect the dynamics and structure at other levels? In social insects, strong hostility towards unrelated individuals can evolve as a kin-selected counter-adaptation to intraspecific social parasitism. This aggression in turn might cause intraspecific competition to predominate over interspecific competition, permitting coexistence with other social insect species. In other words, kin selection—a form of intra-population dynamics—might enhance the species richness of the community, a higher-level structure. The converse effect, from higher to lower levels, might also operate, whereby strong interspecific competition may limit the evolution of selfish individual traits. If the latter effect were to prove more important, it would challenge the common view that intra-population dynamics (via individual or gene selection) is the main driver of evolution.     

Sperm competition,sexual selection and the diverse reproductive biology of Osteoglossiformes

Osteoglossiformes are an order of “bony tongue” fish considered the most primitive living order of teleosts. This review seeks to consolidate known hypotheses and identify gaps in the literature regarding the adaptive significance of diverse reproductive traits and behaviour of osteoglossiforms within the context of sperm competition and the wider lens of sexual selection. Many of the unusual traits observed in osteoglossiforms indicate low levels of sperm competition; most species have unpaired gonads, and mormyroids are the only known vertebrate species with aflagellate sperm. Several osteoglossiform families have reproductive anatomy associated with internal fertilization but perform external fertilization, which may be representative of the evolutionary transition from external to internal fertilization and putative trade-offs between sperm competition and the environment. They also employ every type of parental care seen in vertebrates. Geographically widespread and basally situated within teleosts, osteoglossiforms present an effective study system for understanding how sperm competition and sexual selection have shaped the evolution of teleost reproductive behaviour, sperm and gonad morphology, fertilization strategies, courtship and paternal care, and sexual conflict. The authors suggest that the patterns seen in osteoglossiform reproduction are a microcosm of teleost reproductive diversity, potentially signifying the genetic plasticity that contributed to the adaptive radiation of teleost fishes.     

Effects of within‐colony competition on body size asymmetries and reproductive skew in a social spider

Reproductive partitioning is a key component of social organization in groups of cooperative organisms. In colonies of permanently social spiders of the genus Stegodyphus less than half of the females reproduce, while all females, including nonreproducers, perform suicidal allo‐maternal care. Some theoretical models suggest that reproductive skew is a result of contest competition within colonies, leading to size hierarchies where only the largest females become reproducers. We investigated the effect of competition on within‐group body size variation over six months in S. dumicola, by manipulating food level and colony size. We found no evidence that competition leads to increased size asymmetry within colonies, suggesting that contest competition may not be the proximate explanation for reproductive skew. Within‐colony body size variation was high already in the juvenile stage, and did not increase over the course of the experiment, suggesting that body size variation is shaped at an early stage. This might facilitate task specialization within colonies and ensure colony‐level reproductive output by early allocation of reproductive roles. We suggest that reproductive skew in social spiders may be an adaptation to sociality selected through inclusive fitness benefits of allo‐maternal care as well as colony‐level benefits maximizing colony survival and production.     

Kin selection,kin avoidance and correlated strategies

Summary Kin selection of correlated strategies is examined for both weak and strong altruism under simple haploid inheritance. While kin assortment enhances the range of evolutionary stability for (strongly altruistic) correlated strategies (defined herein), kin avoidance is possible under a weakly altruistic correlated strategy. When social competition induces role assignments of variable fitness, group mates may prefer association with non-relatives. Even when group life is mandatory, an individual may accept the risk of abandonment (and reproductive death) rather then associate with kin: a competitive superior may behave altruistically by permitting competitively inferior kin to emigrate. Thus, kin selection and social competition are not necessarily mutually supportive processes within groups. I conclude by interpreting dominance as a strongly altruistic correlated strategy in two social hymenopteran contexts.     

Costly young and reproductive skew in animal societies

Many recent models of reproductive skew explain subordinatereproduction as a staying incentive offered by dominants, whocan produce more young with a helper present than without. Here,we present a new, alternative explanation for subordinate reproduction,which applies whenever the fitness cost to a parent of producingyoung is an accelerating function of the number produced (ascommonly assumed in optimal clutch size theory). Under these circumstances,a dominant individual may be selected to offer a share of reproductionto a related subordinate, not as an incentive to stay, but becauseadditional offspring that would be expensive for the dominantto produce are cheap for the subordinate. "Beneficial sharing"of this kind is more likely the more closely related the subordinateis to the dominant, so that the model predicts a negative relationshipbetween skew and relatedness. This result runs directly counterto the positive relationship predicted by previous incentive-basedmodels. We explore the interaction of these contrasting effectsby developing an integrated model that allows for both beneficialsharing and staying incentives. When offspring are cheap to produce,this integrated model predicts that the incentive effect will dominate,and skew will increase with relatedness. When young are costly,in contrast, beneficial sharing will be of greater importance,and skew will decrease with relatedness.     

A hypothesis for the evolution of androdioecy: the joint influence of reproductive assurance and local mate competition in a metapopulation

In a subdivided population with recurrent local extinction and re-colonisation, competition amongst related pollen or sperm to fertilise ovules or eggs (‘local mate competition’) is expected to select for female-biased sex allocation. Population turnover should also select against unisexuality in favour of self-fertile cosexuality, because males and females are unable to establish new populations on their own (‘Baker's Law’). Here I argue that androdioecy, a rare breeding system in which males co-occur with hermaphrodites, may evolve in a metapopulation under the joint action of local mate competition and Baker's Law if rates of self-fertilisation decrease with increasing population size. The hypothesis makes several predictions regarding patterns of life-history and sex allocation that are borne out by recent observations of androdioecious species in several unrelated lineages of plants and animals. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparing the consequences of natural selection,adaptive phenotypic plasticity,and matching habitat choice for phenotype–environment matching,population genetic structure,and reproductive isolation in meta‐populations

Organisms commonly experience significant spatiotemporal variation in their environments. In response to such heterogeneity, different mechanisms may act that enhance ecological performance locally. However, depending on the nature of the mechanism involved, the consequences for populations may differ greatly. Building on a previous model that investigated the conditions under which different adaptive mechanisms (co)evolve, this study compares the ecological and evolutionary population consequences of three very different responses to environmental heterogeneity: matching habitat choice (directed gene flow), adaptive plasticity (associated with random gene flow), and divergent natural selection. Using individual‐based simulations, we show that matching habitat choice can have a greater adaptive potential than plasticity or natural selection: it allows for local adaptation while protecting genetic polymorphism despite global mating or strong environmental changes. Our simulations further reveal that increasing environmental fluctuations and unpredictability generally favor the emergence of specialist genotypes but that matching habitat choice is better at preventing local maladaptation by individuals. This confirms that matching habitat choice can speed up the genetic divergence among populations, cause indirect assortative mating via spatial clustering, and hence even facilitate sympatric speciation. This study highlights the potential importance of directed dispersal in local adaptation and speciation, stresses the difficulty of deriving its operation from nonexperimental observational data alone, and helps define a set of ecological conditions which should favor its emergence and subsequent detection in nature.     

Habitat selection in rock pool corixids: the effect of local density on dispersal

The corixid species breeding in temporary rock pools of Baltic archipelago live in a highly fragmented and unpredictable habitat. Shallow rock pools can dry out and be refilled repeatedly during a summer causing high mortality of immatures. In deeper pools, young nymphs face intense competition by older stages including cannibalism. The adult corixids move frequently between rock pools and are thus able to use currently available habitat for reproduction. In this dispersal behaviour, the ability to assess the local population density and hence select the more suitable low density patches would be advantageous. We studied the effect of local population density on the frequency of dispersive flights of Arctocorisa carinata (Sahlberg) and Callicorixa producta (Reuter) experimentally, using rock pools from which nymphs of both species were removed. The dispersal rates of marked C. producta adults were significantly lower from experimental rock pools than from normal density controls, leading to a concentration of C. producta adults in the experimental rock pools. Indications of immigration rate differences between the experimental and control pools were also observed. No clear differences were found in the superior competitor A. carinata.     

Small fish,large river: Surprisingly minimal genetic structure in a dispersal‐limited,habitat specialist fish

Genetic connectivity is expected to be lower in species with limited dispersal ability and a high degree of habitat specialization (intrinsic factors). Also, gene flow is predicted to be limited by habitat conditions such as physical barriers and geographic distance (extrinsic factors). We investigated the effects of distance, intervening pools, and rapids on gene flow in a species, the Tuxedo Darter (Etheostoma lemniscatum), a habitat specialist that is presumed to be dispersal‐limited. We predicted that the interplay between these intrinsic and extrinsic factors would limit dispersal and lead to genetic structure even at the small spatial scale of the species range (a 38.6 km river reach). The simple linear distribution of E. lemniscatum allowed for an ideal test of how these factors acted on gene flow and allowed us to test expectations (e.g., isolation‐by‐distance) of linearly distributed species. Using 20 microsatellites from 163 individuals collected from 18 habitat patches, we observed low levels of genetic structure that were related to geographic distance and rapids, though these factors were not barriers to gene flow. Pools separating habitat patches did not contribute to any observed genetic structure. Overall, E. lemniscatum maintains gene flow across its range and is comprised of a single population. Due to the linear distribution of the species, a stepping‐stone model of dispersal best explains the maintenance of gene flow across its small range. In general, our observation of higher‐than‐expected connectivity likely stems from an adaptation to disperse due to temporally unstable and patchy habitat.     

改良拔河针法对膝关节骨性关节炎患者远期关节功能和生活质量的影响

摘要 目的：探讨改良拔河针法对膝关节骨性关节炎患者远期关节功能和生活质量的影响，以期为膝关节骨性关节炎患者的治疗提供更加合理方案。方法：选取2016年3月至2019年10月深圳市中西医结合医院收治的60例膝关节骨性关节炎患者为研究对象。采用系统随机化法分为常规针刺组、改良拔河针法组各30例。常规针刺组采用常规针刺治疗，改良拔河针法组采用改良拔河针法治疗。分别于疗程结束时比较两组临床有效率，于治疗前、治疗后及治疗后3个月比较两组患者疼痛评级指数（PRI）评分、Oswestry功能障碍指数问卷表（ODI）评分及简明健康状况调查量表（SF-36）评分。结果：改良拔河针法组临床有效率86.67%，略高于常规针刺组的80.00%，但组间比较差异无统计学意义（P＞0.05）。组内比较显示，两组患者治疗后、治疗后3个月的PRI评分、ODI评分及SF-36评分较治疗前均降低（P＜0.05），且治疗后3个月的PRI评分、ODI评分及SF-36评分低于治疗后（P＜0.05）。组间比较显示，两组患者治疗前、治疗后PRI评分、ODI评分及SF-36评分比较差异均无统计学意义（P＞0.05），但改良拔河针法组治疗后3个月的ODI评分及SF-36评分优于常规针刺组（P＜0.05）。结论：改良拔河针法治疗膝关节骨性关节炎患者可获得普通针刺法相当的临床效果，且在改善患者远期关节功能和生活质量方面效果更佳，值得临床推广应用。     

Origins of altruism diversity I: the diverse ecological roles of altruistic strategies and their evolutionary responses to local competition

Nature abounds with a rich variety of altruistic strategies, including public resource enhancement, resource provisioning, communal foraging, alarm calling, and nest defense. Yet, despite their vastly different ecological roles, current theory typically treats diverse altruistic traits as being favored under the same general conditions. Here, we introduce greater ecological realism into social evolution theory and find evidence of at least four distinct modes of altruism. Contrary to existing theory, we find that altruistic traits contributing to "resource-enhancement" (e.g., siderophore production, provisioning, agriculture) and "resource-efficiency" (e.g., pack hunting, communication) are most strongly favored when there is strong local competition. These resource-based modes of helping are "K-strategies" that increase a social group's growth yield, and should characterize species with scarce resources and/or high local crowding caused by low mortality, high fecundity, and/or mortality occurring late in the process of resource-acquisition. The opposite conditions, namely weak local competition (abundant resource, low crowding), favor survival (e.g., nest defense) and fecundity (e.g., nurse workers) altruism, which are "r-strategies" that increase a social group's growth rate. We find that survival altruism is uniquely favored by a novel evolutionary force that we call "sunk cost selection." Sunk cost selection favors helping that prevents resources from being wasted on individuals destined to die before reproduction. Our results contribute to explaining the observed natural diversity of altruistic strategies, reveal the necessary connection between the evolution and the ecology of sociality, and correct the widespread but inaccurate view that local competition uniformly impedes the evolution of altruism.     

Kin selection, quorum sensing and virulence in pathogenic bacteria

Bacterial growth and virulence often depends upon the cooperative release of extracellular factors excreted in response to quorum sensing (QS). We carried out an in vivo selection experiment in mice to examine how QS evolves in response to variation in relatedness (strain diversity), and the consequences for virulence. We started our experiment with two bacterial strains: a wild-type that both produces and responds to QS signal molecules, and a lasR (signal-blind) mutant that does not release extracellular factors in response to signal. We found that: (i) QS leads to greater growth within hosts; (ii) high relatedness favours the QS wild-type; and (iii) low relatedness favours the lasR mutant. Relatedness matters in our experiment because, at relatively low relatedness, the lasR mutant is able to exploit the extracellular factors produced by the cells that respond to QS, and hence increase in frequency. Furthermore, our results suggest that because a higher relatedness favours cooperative QS, and hence leads to higher growth, this will also lead to a higher virulence, giving a relationship between relatedness and virulence that is in the opposite direction to that usually predicted by virulence theory.     

Male reproductive skew, paternal relatedness, and female social relationships

Female social relationships among primates are thought to be shaped by socio-ecological factors and phylogenetic constraints. We suggest that patterns of paternal relatedness among females influence measures of social tolerance that have been used to classify species into different social relationship categories. As kin support and kin preference have only been measured for matrilineal kin and related individuals exchange less aggression and have a higher conciliatory tendency, the observed low nepotism levels and high tolerance levels may be an artifact of hidden paternal relatedness among the nonkin category. Using comparative data on macaques, we investigate this hypothesis using male reproductive skew as a proxy for paternal relatedness. Within the limitations of the study we show that populations classified as being less nepotistic, and more tolerant exhibit higher levels of reproductive skew. This first result and the reasoning behind may motivate future students of social relationships to take paternal relatedness into consideration. Potential implications of this finding if repeated with larger samples include that variation in aspects of macaque social relationships may be explained without considering phylogeny or the strength of between-group contest competition for food.     

Kin and kinship psychology both influence cooperative coordination in Yasawa,Fiji

Genes shared through common ancestry are among the oldest social bonds. Despite these ancient roots, humans often co-opt the psychology of genetic relatedness and extend it to genetically unrelated others through culturally-acquired kinship systems. We investigate how genealogical relatedness and kinship norms might mutually support or oppose each other within a known kin network in Yasawa, Fiji. Yasawans' reliance on intensive, kin-based cooperation for daily life makes Yasawan kinship an interesting test case to compare the effects of genealogy and kinship norms. Confirming qualitative ethnographic claims, we find that Yasawan kin terms can be described in two dimensions of respect/closeness and joking/authority. Individual players use different strategies for genealogical relatives and non-relatives by making economic game choices that are increasingly beneficial to partners who share a higher percentage of genes through common ancestry. However, pairs of players are most successful in coordinating their game choices despite conflicting self-interests based upon kinship norms relevant to hierarchy. Thus, while genealogical relatedness may boost generosity, the extra behavioral structuring from kinship norms facilitates more productive but difficult coordinated action even when communication is not possible.