Inclusive fitness theory and eusociality

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Nowak et al. argue that inclusive fitness theory has been of little value in explaining the natural world, and that it has led to negligible progress in explaining the evolution of eusociality. However, we believe that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature. We will focus our comments on three general issues.     

In defence of inclusive fitness theory

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Arguably the defining characteristic of the scientific process is its capacity for self-criticism and correction. Nowak et al. challenge proposed connections between relatedness and the evolution of eusociality, suggest instead that defensible nests and "spring-loaded" traits are key, and present alternative modelling approaches. They then dismiss the utility of Hamilton's insight that relatedness has a profound evolutionary effect, formalized in his widely accepted inclusive fitness theory as Hamilton's rule ("Rise and fall of inclusive fitness theory"). However, we believe that Nowak et al. fail to make their case for logical, theoretical and empirical reasons.     

Kin selection and eusociality

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Hamilton described a selective process in which individuals affect kin (kin selection), developed a novel modelling strategy for it (inclusive fitness), and derived a rule to describe it (Hamilton's rule). Nowak et al. assert that inclusive fitness is not the best modelling strategy, and also that its production has been "meagre". The former may be debated by theoreticians, but the latter is simply incorrect. There is abundant evidence to demonstrate that inclusive fitness, kin selection and Hamilton's rule have been extraordinarily productive for understanding the evolution of sociality.     

Only full-sibling families evolved eusociality

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. The paper by Nowak et al. has the evolution of eusociality as its title, but it is mostly about something else. It argues against inclusive fitness theory and offers an alternative modelling approach that is claimed to be more fundamental and general, but which, we believe, has no practical biological meaning for the evolution of eusociality. Nowak et al. overlook the robust empirical observation that eusociality has only arisen in clades where mothers are associated with their full-sibling offspring; that is, in families where the average relatedness of offspring to siblings is as high as to their own offspring, independent of population structure or ploidy. We believe that this omission makes the paper largely irrelevant for understanding the evolution of eusociality.     

Fine-scale genetic structuring on Manacus manacus leks

Leks have traditionally been considered as arenas where males compete to attract females and secure matings. Thus, direct fitness benefits mediated through competition between males to fertilize females have been considered to be the primary force driving the evolution of lekking behaviour. Inclusive fitness benefits mediated through kin selection may also be involved in lek formation and evolution, but to date this theory has been largely ignored. According to kin-selection theory, both reproducing and non-reproducing males may gain indirect inclusive fitness benefits. If females are attracted to larger leks, non-reproducing males add attractiveness to a lek, and therefore, in a genetically structured population, boost the reproductive success of kin. Theory predicts that the attractiveness of leks is plastic, and that males establish themselves on a lek in which the top male, in terms of reproductive success, is a close relative. Here we show that in white-bearded manakins (Manacus manacus), for which larger leks are more attractive to females and so secure the maximum number of matings, there is extraordinary fine-scale genetic structure, with leks being composed of clusters of related kin. We propose that males establish themselves where they find relatives to such an extent that they form groups within leks, and that such behaviour is consistent with kin-selection theory to maximize reproductive success of the group.     

合作进化的研究进展

阐述了合作的进化对于理解生物（包括人类自身）组织行为的重要性．合作的进化描述了生物行为策略的选择过程,互相剥夺还是互相合作．在理论模型中则取决于作用双方再次相遇的概率（即未来贴现）、环境噪声的大小、人口的结构特征、对策的发生机制等因素．较优的策略能在长期内获得较高的支付值,表征了个体对环境的适应性,体现了自然选择的结果．同时探讨了进化策略的稳定性,以及由Ｎｏｗａｋ 等引入的反映合作疆域之争的空间对策问题     

Life-history trade-offs favour the evolution of animal personalities

In recent years evidence has been accumulating that personalities are not only found in humans but also in a wide range of other animal species. Individuals differ consistently in their behavioural tendencies and the behaviour in one context is correlated with the behaviour in multiple other contexts. From an adaptive perspective, the evolution of animal personalities is still a mystery, because a more flexible structure of behaviour should provide a selective advantage. Accordingly, many researchers view personalities as resulting from constraints imposed by the architecture of behaviour (but see ref. 12). In contrast, we show here that animal personalities can be given an adaptive explanation. Our argument is based on the insight that the trade-off between current and future reproduction often results in polymorphic populations in which some individuals put more emphasis on future fitness returns than others. Life-history theory predicts that such differences in fitness expectations should result in systematic differences in risk-taking behaviour. Individuals with high future expectations (who have much to lose) should be more risk-averse than individuals with low expectations. This applies to all kinds of risky situations, so individuals should consistently differ in their behaviour. By means of an evolutionary model we demonstrate that this basic principle results in the evolution of animal personalities. It simultaneously explains the coexistence of behavioural types, the consistency of behaviour through time and the structure of behavioural correlations across contexts. Moreover, it explains the common finding that explorative behaviour and risk-related traits like boldness and aggressiveness are common characteristics of animal personalities.     

Stepwise evolution of stable sociality in primates

Although much attention has been focused on explaining and describing the diversity of social grouping patterns among primates, less effort has been devoted to understanding the evolutionary history of social living. This is partly because social behaviours do not fossilize, making it difficult to infer changes over evolutionary time. However, primate social behaviour shows strong evidence for phylogenetic inertia, permitting the use of Bayesian comparative methods to infer changes in social behaviour through time, thereby allowing us to evaluate alternative models of social evolution. Here we present a model of primate social evolution, whereby sociality progresses from solitary foraging individuals directly to large multi-male/multi-female aggregations (approximately 52 million years (Myr) ago), with pair-living (approximately 16?Myr ago) or single-male harem systems (approximately 16?Myr ago) derivative from this second stage. This model fits the data significantly better than the two widely accepted alternatives (an unstructured model implied by the socioecological hypothesis or a model that allows linear stepwise changes in social complexity through time). We also find strong support for the co-evolution of social living with a change from nocturnal to diurnal activity patterns, but not with sex-biased dispersal. This supports suggestions that social living may arise because of increased predation risk associated with diurnal activity. Sociality based on loose aggregation is followed by a second shift to stable or bonded groups. This structuring facilitates the evolution of cooperative behaviours and may provide the scaffold for other distinctive anthropoid traits including coalition formation, cooperative resource defence and large brains.     

Complex social behaviour derived from maternal reproductive traits

A fundamental goal of sociobiology is to explain how complex social behaviour evolves, especially in social insects, the exemplars of social living. Although still the subject of much controversy, recent theoretical explanations have focused on the evolutionary origins of worker behaviour (assistance from daughters that remain in the nest and help their mother to reproduce) through expression of maternal care behaviour towards siblings. A key prediction of this evolutionary model is that traits involved in maternal care have been co-opted through heterochronous expression of maternal genes to result in sib-care, the hallmark of highly evolved social life in insects. A coupling of maternal behaviour to reproductive status evolved in solitary insects, and was a ready substrate for the evolution of worker-containing societies. Here we show that division of foraging labour among worker honey bees (Apis mellifera) is linked to the reproductive status of facultatively sterile females. We thereby identify the evolutionary origin of a widely expressed social-insect behavioural syndrome, and provide a direct demonstration of how variation in maternal reproductive traits gives rise to complex social behaviour in non-reproductive helpers.     

Evolution of cooperation in a finite homogeneous graph

Recent theoretical studies of selection in finite structured populations have worked with one of two measures of selective advantage of an allele: fixation probability and inclusive fitness. Each approach has its own analytical strengths, but given certain assumptions they provide equivalent results. In most instances the structure of the population can be specified by a network of nodes connected by edges (that is, a graph), and much of the work here has focused on a continuous-time model of evolution, first described by ref. 11. Working in this context, we provide an inclusive fitness analysis to derive a surprisingly simple analytical condition for the selective advantage of a cooperative allele in any graph for which the structure satisfies a general symmetry condition ('bi-transitivity'). Our results hold for a broad class of population structures, including most of those analysed previously, as well as some for which a direct calculation of fixation probability has appeared intractable. Notably, under some forms of population regulation, the ability of a cooperative allele to invade is seen to be independent of the nature of population structure (and in particular of how game partnerships are specified) and is identical to that for an unstructured population. For other types of population regulation our results reveal that cooperation can invade if players choose partners along relatively 'high-weight' edges.     

进化计算中的精英个体产生策略

初始种群是影响基因表达式编程算法（GEP）的重要因素之一。提出了精英个体产生策略,产生具有较高的适应度的精英个体,使种群从一个较高的基础上开始进化,从而提高种群的进化效率。在此基础上,提出了综合精英个体产生策略和基因空间均匀分布策略优点的综合种群产生算法。实验表明,精英个体产生策略可以提高进化效率17%,综合算法可以更加有效地提高系统的进化效率。以上算法的思想还可以应用于其它进化计算中。     

Mate fidelity and intra-lineage polygyny in greater horseshoe bats

Mating strategies that lead to increased kinship within socially cooperative groups may offer inclusive fitness benefits to individuals, but can also result in higher levels of inbreeding. Here we show in a sexually segregated bat species that females avoid this conflict through two mating behaviours. First, most females revisit and breed with specific, individual males across years, so that their single offspring born in different years are full siblings. Second, relatives in the maternal line, including mothers and daughters, share breeding partners (intra-lineage polygyny) more often than expected by chance. Although these behaviours increased levels of co-ancestry among colony members, there was no concomitant rise in inbreeding. We suggest that when females engage in mate fidelity and intra-lineage polygyny, kin ties among female roost mates will be strengthened, thereby potentially contributing to social group cohesiveness. Our findings reveal the hidden complexity that can underlie polygynous breeding, and highlight a new potential route by which female mate choice could influence social evolution.     

适用于大数据的遗传优化算法研究

针对传统遗传算法对高维度数据或大数据易陷入局部最优的问题,提出了一种基于网格小生境与分级多种群共同演化的增强遗传算法.首先,采用基于网格的小生境算法建立主种群,主种群独立进化,将低适应度的样本迁移至子种群中.按照低适应度样本的适应度范围建立分级的子种群结构,各子种群内的样本独立演化,低适应度子种群的样本可进化并迁移至高适应度种群或返回主种群,从而防止具有一定竞争力的样本过早死亡.对比实验结果表明,本算法对高维度数据具有较好的优化效果,优于同类型遗传算法.     

法学研究的新范式——进化生物学范式及其展望

进化生物学已对几乎所有社会科学产生影响，并成为经济学、心理学和人类学等学科的主导研究范式之一，但在法学研究中仍未取得与之相称的进展。进化生物学既研究有机体进化也研究文化进化，法律是一种文化现象并有其进化上的根源，因而在法学研究中引入进化生物学范式就成为可能。     

Protein dispensability and rate of evolution.

If protein evolution is due in large part to slightly deleterious amino acid substitutions, then the rate of evolution should be greater in proteins that contribute less to individual fitness. The rationale for this prediction is that relatively dispensable proteins should be subject to weaker purifying selection, and should therefore accumulate mildly deleterious substitutions more rapidly. Although this argument was presented over twenty years ago, and is fundamental to many applications of evolutionary theory, the prediction has proved difficult to confirm. In fact, a recent study showed that essential mouse genes do not evolve more slowly than non-essential ones. Thus, although a variety of factors influencing the rate of protein evolution have been supported by extensive sequence analysis, the relationship between protein dispensability and evolutionary rate has remained unconfirmed. Here we use the results from a highly parallel growth assay of single gene deletions in yeast to assess protein dispensability, which we relate to evolutionary rate estimates that are based on comparisons of sequences drawn from twenty-one fully annotated genomes. Our analysis reveals a highly significant relationship between protein dispensability and evolutionary rate, and explains why this relationship is not detectable by categorical comparison of essential versus non-essential proteins. The relationship is highly conserved, so that protein dispensability in yeast is also predictive of evolutionary rate in a nematode worm.     

How did brains evolve?

Three reports on mammalian brain evolution analyse the same comparative data on brain component volumes but come to partially conflicting conclusions. Clark et al. conclude from their analysis of volumetric brain proportions ("cerebro-types") that cerebellum size is invariant across mammalian taxonomic groups, the neocortex and cerebellum do not co-vary in size (in contradiction to ref. 1), and cerebrotype-based measures identify directional changes in brain architecture. Here I provide evidence that calls each of these conclusions into question. The failure of the cerebrotype measure to identify species differences in brain architecture that are independent of gross brain size undermines the proposal by Clark et al. that it could be useful for detecting evolutionary patterns and phylogenetic relationships.     

Do female hyaenas choose mates based on tenure?

In their investigation into whether female mate-choice drives male dispersal, H?ner et al. argue that female spotted hyaenas (Crocuta crocuta) prefer mates whose tenure in the social group is less than the females' age, to avoid paternal incest, and suggest that male dispersal reflects this preference. However, we are not persuaded that females choose mates on the basis of tenure because H?ner et al. overlook the alternative hypothesis that dispersal status itself is important in female mate-choice, such that females prefer immigrants over natal males. Like mate-choice based on tenure, choice based on dispersal status reduces the risk of incest.     

Biodiversity conservation: effects of changes in climate and land use

Thomas et al. argue, contrary to Sala et al. that climate change poses an equal or greater threat to global biodiversity than land-use change. We contest this claim, however, on the grounds that Thomas et al. incorrectly apply species-area relationships.     

贵州遵义下寒武统牛蹄塘组生物群中的海绵化石

遵义松林中南村下寒武统牛蹄塘组中下部,特别是下部黑色泥岩中产有大量的海绵骨针,形态多样,初步研究有多达10属以上的海绵动物,与澄江生物群的海绵相比,牛蹄塘组的海绵动物化石其组合特征以六射海绵动物为主,它的发现增添了寒武纪生物组成的信息,为海绵动物从晚新元古代的发生以至穿越寒武纪的演化过程及寒武纪生物大爆发的定性,量度提供了重要信息及资料,描述了4属2个种。1个相似种和1个未定种。Hyalosinica sp.;Saetaspongia densa Mehl et Reitner,1993;Leptomitus teretiusculus CHen,Hou et Lu.1989;Crumillospongia cf.biporosa Righy,1986,它们分属于海绵动物的六射海绵纲和普通海绵纲。     

Learned kin recognition cues in a social bird

In many cooperatively breeding birds, kin selection has an important role in the evolution and maintenance of social behaviour, and 'helpers' can maximize indirect fitness gains by preferentially allocating care to close relatives. Although there is evidence for kin-biased helping behaviour in several species, the mechanism of kin recognition underlying this behaviour is poorly understood. Vocalizations are the most commonly used cues in avian recognition systems, but the effectiveness of vocal signals as reliable recognition cues must depend on how they are acquired. However, there have been no experimental studies of the development of vocal recognition cues in cooperative birds; indeed, the ontogeny of all bird vocalizations other than song is poorly known in any species. Here, we show that cooperatively breeding long-tailed tits (Aegithalos caudatus) can discriminate between kin and non-kin according to the individual-specific characteristics of contact calls, and show experimentally that individuals learn these calls from provisioning adults during the nestling period. Finally, we show that the pattern of cooperative behaviour in this species is consistent with the use of recognition cues learned through association.