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.     

Inclusive fitness in evolution

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. For over fifty years, the evolution of social behaviour has been guided by the concept of inclusive fitness as a measure of evolutionary success. Nowak et al. argue that inclusive fitness should be abandoned. In so doing, however, they misrepresent the role that inclusive fitness has played in the theory of social evolution by which understanding social behaviour in a variety of disciplines has developed and flourished. By discarding inclusive fitness on the basis of its limitations, they create a conceptual tension which, we argue, is unnecessary, and potentially dangerous for evolutionary biology.     

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.     

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.     

Kin selection and cooperative courtship in wild turkeys

In the few species of birds in which males form display partnerships to attract females, one male secures most or all of the copulations. This leads to the question of why subordinate males help in the absence of observable reproductive benefits. Hamilton's concept of kin selection, whereby individuals can benefit indirectly by helping a relative, was a crucial breakthrough for understanding apparently altruistic systems. However in the only direct test of kin selection in coordinated display partnerships, partners were unrelated, discounting kin selection as an explanation for the evolution of cooperation. Here I show, using genetic measures of relatedness and reproductive success, that kin selection can explain the evolution of cooperative courtship in wild turkeys. Subordinate (helper) males do not themselves reproduce, but their indirect fitness as calculated by Hamilton's rule more than offsets the cost of helping. This result confirms a textbook example of kin selection that until now has been controversial and also extends recent findings of male relatedness on avian leks by quantifying the kin-selected benefits gained by non-reproducing males.     

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.     

Testing Hamilton's rule with competition between relatives

Hamilton's theory of kin selection suggests that individuals should show less aggression, and more altruism, towards closer kin. Recent theoretical work has, however, suggested that competition between relatives can counteract kin selection for altruism. Unfortunately, factors that tend to increase the average relatedness of interacting individuals--such as limited dispersal--also tend to increase the amount of competition between relatives. Therefore, in most natural systems, the conflicting influences of increased competition and increased relatedness are confounded, limiting attempts to test theory. Fig wasp taxa exhibit varying levels of aggression among non-dispersing males that show a range of average relatedness levels. Thus, across species, the effects of relatedness and competition between relatives can be separated. Here we report that--contrary to Hamilton's original prediction but in agreement with recent theory--the level of fighting between males shows no correlation with the estimated relatedness of interacting males, but is negatively correlated with future mating opportunities.     

Communal nesting patterns in mice implicate MHC genes in kin recognition.

House mice (Mus musculus domesticus) form communal nests and appear to nurse each other's pups indiscriminately. Communal nesting probably functions to reduce infanticide, but it also makes females vulnerable to exploitation if nursing partners fail to provide their fair share of care. Kinship theory predicts that females will preferentially form communal nests with relatives to minimize exploitation and further increase inclusive fitness. Here we provide evidence from seminatural populations that females prefer communal nesting partners that share allelic forms of major histocompatibility complex genes. Such behaviour would lead to the selection of close relatives as communal nesting partners. Although criteria for the demonstration of kin recognition are currently embroiled in controversy, this is the first vertebrate study to meet Grafen's restrictive requirements: discrimination is based on genetic similarity at highly polymorphic loci, incidental correlations due to relatedness are experimentally controlled, and strong reasons exist for expecting the assayed behaviour to be kin-selected.     

Genetic relatedness predicts South African migrant workers' remittances to their families

Inclusive fitness models predict many commonly observed behaviours: among humans, studies of within-household violence, the allocation of food and child care find that people favour those to whom they are more closely related. In some cases however, kin-altruism effects appear to be modest. Do individuals favour kin to the extent that kin-altruism models predict? Data on remittances sent by South African migrant workers to their rural households of origin allow an explicit test, to our knowledge the first of its kind for humans. Using estimates of the fitness benefits and costs associated with the remittance, the genetic relatedness of the migrant to the beneficiaries of the transfer, and their age- and sex-specific reproductive values, we estimate the level of remittance that maximizes the migrant worker's inclusive fitness. This is a much better predictor of observed remittances than is average relatedness, even when we take account (by means of a multiple regression) of covarying influences on the level of remittance. But the effect is modest: less than a third of the observed level of remittances can be explained by our kin-altruism model.     

“阳光体育运动”背景下高校开展健美操运动存在问题分析及对策研究

"阳光体育运动"是新时期加强青少年体育、增强青少年体质的战略举措,是各级学校贯彻"健康第一"指导思想和形成"终身体育"行为习惯的具体体现.在此背景下,高校在健美操教学与开展上存在观念落后、忽视了对学生运动兴趣的培养、教学结构和内容缺乏系统性等问题,并提出相应的改进措施.     

Short-latency drinking and increased Na appetite after intracerebral microinjections of NGF in rats

NERVE GROWTH FACTOR (NGF) is a polypeptide trophic factor for peripheral sympathetic and sensory neurones(1,2). Apparent NGF(3-5) and NGF receptors(6,7) have also been identified in the brain, and intracerebral administration of NGF in the adult rat produces marked biochemical(8) and morphologica(9,10) changes in brain tissue. These findings, taken together with the observations that central injections of NGF facilitate behavioural recovery from brain damage(11,12), indicate that this polypeptide may have an important role in brain function. It has been observed that rats given intraventricular injections of up to 2.3 microg NGF drink copiously (M.E.L. and G. Guroff, unpublished observations). Perkins et al.(13) reported that diencephalic application of crystalline NGF (1-15 microg) resulted in an intense polydipsia. The present report confirms the observations of M.E.L. and Guroff, and extends the findings of Perkins et al.(13) by using solutions of NGF instead of crystals. It also describes for the first time a second phenomenon produced by intracranial administration of NGF, namely an intense appetite for aversive concentrations of sodium solutions.     

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.     

Experimental quest for orbital waves

One challenge in condensed-matter physics is the experimental confirmation of a new kind of elementary excitation orbital waves, or orbitons, which are predicted to exist in an orbitally ordered state. Saitoh et al. have observed three peaks at 160, 144 and 126 meV in the Raman scattering of orbitally ordered lanthanum manganate (LaMnO(3)), and interpret these as evidence of orbitons. However, we find similar peaks in the optical conductivity, sigma(omega), of LaMnO(3) and point out that the direct observation of orbitons in sigma(omega) is prohibited by a selection rule. This suggests that the Raman peaks observed by Saitoh et al. arise from multiphonons, and that the existence of orbitons has yet to be experimentally confirmed.     

BBM方程的Hamilton表示

研究了水波动力学中重要规则长波方程之一的BBM方程的Hamilton表示.运用Olver研究该类方程的守恒律来定义能量函数E(u),推导出一种新型的变分原理,并利用所得到的变分原理导出BBM方程的Hamilton表示.同时,利用待定泛函形式的变分原理导出另一种形式的BBM方程的Hamilton表示.两种形式的Hamilton表示均可引进辛差分格式,进行数值解计算.     

Configuration of the magnetic field and reconstruction of Pangaea in the Permian period

The virtual geomagnetic poles of Laurasia and Gondwanaland in the Carboniferous and Permian periods diverge significantly when these continents are reassembled according to the fit calculated by Bullard et al. Two interpretations have been offered: Briden et al. explain these divergences by a magnetic field configuration very different from that of a geocentric axial dipole; Irving (and private communication), Van der Voo and French(4) suggest a different reconstruction and it is shown here that these two interpretations are not incompatible and that the first may help the second.     

基于遗传编程的分等级规则挖掘模型

根据分等级规则挖掘的具体要求,结合遗传编程的特点建立了一种新型的规则挖掘模型。对规则的编码方式和遗传算子进行了详细的设计,构造了满足遗传编程挖掘的树型染色体规则结构,并综合考虑支持度、可信度和相关度3个指标和等级系数等约束因素来设计适应度函数。最后通过实例对基于遗传编程的分等级规则挖掘模型的可靠性进行了验证。     

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.     

An evolutionarily stable strategy approach to indiscriminate spite

AN individual behaves spitefully when it harms itself in order to harm another individual more(1). Hamilton(1,2) predicted that spite may evolve if it is expressed only in those encounters that occur between individuals of less than average relatedness. More recently Verner(3) suggested that territory size may become super-optimal because of a selective advantage arising from the spiteful exclusion of others from limited resources. His model is essentially different from Hamilton's in that spite is directed at individuals indiscriminately with respect to relatedness. Recently Rothstein(4) has shown analytically that the initial spread of spiteful traits will be very slow in all but the smallest populations. He also argued verbally that indiscriminate spite can never be evolutionarily stable even if it should spread (see also Davies(5)). The question of evolutionary stability is clearly important, but its resolution requires an analytical approach. We report here an approach based on Maynard Smith's(6) concept of the evolutionarily stable strategy (ESS), a strategy which, when common, does better than any alternative strategy played by a rare mutant. We show that spite can be an ESS, but that the magnitude of spite will be small in large populations.     

A new approach based on PSO algorithm to find good computational encoding sequences

Computational encoding DNA sequence design is one of the most important steps in molecular computation. A lot of research work has been done to design reliable sequence library. A revised method based on the support system developed by Tanaka et al. is proposed here with different criteria to construct fitness function. Then we adapt particle swarm optimization (PSO) algorithm to our encoding problem. By using the new algorithm, a set of sequences with good quality is generated. The result also shows that our PSO-based approach could rapidly converge at the minimum level for an output of the simulation model. The celerity of the algorithm fits our requirements.     

一种基于ELECTRE法的多目标优化进化算法

将多目标属性决策方法中的ELECTRE法引入到多目标优化进化算法中,提出了一种新的多目标优化算法.采用辅助群体来存储进化过程中的非劣个体,并且采用与SPEA-Ⅱ相同的适应值分配策略来保证解的良好分布性.此外,构造出一种新的超序关系对个体进行排序,证明了该超序关系比Pareto优劣关系弱,利用此超序关系,能增强进化过程中的选择压,加快收敛速度.数据实验结果表明,该算法能很好地收敛到Pareto最优,有效地保持解的多样性.