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In many cooperative societies, including our own, helpers assist with the post-natal care of breeders' young and may thereby benefit the post-natal development of offspring. Here, we present evidence of a novel mechanism by which such post-natal helping could also have beneficial effects on pre-natal development: By lightening post-natal maternal workloads, helpers may allow mothers to increase their pre-natal investment per offspring. We present the findings of a decade-long study of cooperatively breeding white-browed sparrow-weaver, Plocepasser mahali, societies. Within each social group, reproduction is monopolized by a dominant breeding pair, and non-breeding helpers assist with nestling feeding. Using a within-mother reaction norm approach to formally identify maternal plasticity, we demonstrate that when mothers have more female helpers, they decrease their own post-natal investment per offspring (feed their nestlings at lower rates) but increase their pre-natal investment per offspring (lay larger eggs, which yield heavier hatchlings). That these plastic maternal responses are predicted by female helper number, and not male helper number, implicates the availability of post-natal helping per se as the likely driver (rather than correlated effects of group size), because female helpers feed nestlings at substantially higher rates than males. We term this novel maternal strategy "maternal front-loading" and hypothesize that the expected availability of post-natal help either allows or incentivizes helped mothers to focus maternal investment on the pre-natal phase, to which helpers cannot contribute directly. The potential for post-natal helping to promote pre-natal development further complicates attempts to identify and quantify the fitness consequences of helping. In many cooperative societies, including our own, helpers assist with the post-natal care of breeders' young, and may thereby benefit the post-natal development of offspring. This study shows that mothers in a wild cooperatively breeding bird lay larger eggs when they will have more help with nestling care. This strategy may allow helped mothers to focus maternal investment on the pre-natal phase, to which helpers cannot contribute directly. [ABSTRACT FROM AUTHOR]

What explains the outcomes of chance processes? We claim that their setups do. Chances, we think, mediate these explanations of outcome by setup but do not feature in them. Facts about chances do feature in explanations of a different kind: higher-order explanations, which explain how and why setups explain their outcomes. In this paper, we elucidate this 'mediator view' of chancy explanation and defend it from a series of objections. We then show how it changes the playing field in four metaphysical disputes concerning chance. First, it makes it more plausible that even low chances can have explanatory power. Second, it undercuts a circularity objection against reductionist theories of chance. Third, it redirects the debate about a prominent argument against epistemic theories of chance. Finally, it sheds light on potential chancy explanations of the Universe's origin. [ABSTRACT FROM AUTHOR]

Sex differences in lifespan remain an intriguing puzzle in evolutionary biology. While explanations range from sex differences in selection to sex differences in the expression of recessive lifespan‐altering mutations (via X‐linkage), little consensus has been reached. One unresolved issue is the extent to which genetic influences on lifespan dimorphism are modulated by the environment. For example, studies have shown that sex differences in lifespan can either increase or decrease depending upon the social environment. Here, we took an experimental approach, manipulating multiple axes of the social environment across inbred long‐ and short‐lived genotypes and their reciprocal F1s in the fly Drosophila serrata. Our results reveal strong genetic effects and subtle yet significant genotype‐by‐environment interactions for male and female lifespan, specifically due to both population density and mating status. Further, our data do not support the idea that unconditional expression of deleterious X‐linked recessive alleles in heterogametic males accounts for lower male lifespan. [ABSTRACT FROM AUTHOR]

This article explores three ways in which physics may involve counterpossible reasoning. The first way arises when evaluating false theories: to say what the world would be like if the theory were true, we need to evaluate counterfactuals with physically impossible antecedents. The second way relates to the role of counterfactuals in characterizing causal structure: to say what causes what in physics, we need to make reference to physically impossible scenarios. The third way is novel: to model metaphysical dependence in physics, we need to consider counterfactual consequences of metaphysical impossibilities. Physics accordingly bears substantial and surprising counterpossible commitments. [ABSTRACT FROM AUTHOR]

Exposure to and potential to engage in music is now very different for children and young people from more middle‐class and affluent backgrounds compared to those from working‐class or poor households. Middle‐class children and young people have access to opportunities to learn an instrument and engage in performance, predominantly for intrinsic reward and on occasion as the basis of a future career. Children and young people from more working class or poor backgrounds are more likely to be excluded from these opportunities and are less likely to take advantage of opportunities within the wider world of music industries. This research draws on in depth case studies of three local authorities in Scotland to reveal the ways in which local provision for music education is largely being reshaped by economic factors and restraints. Drawing on aspects of critical theory it discusses the ways in which economic‐based decisions are interacting with the dominant social and cultural values to influence provision. It makes the argument that in an era of austerity the social and cultural values of the middle class are dominant and effectively increasing inequality in access to music. Children and young people from poor or working class households are increasingly excluded from opportunities to engage with formal music provision. [ABSTRACT FROM AUTHOR]

Theoretical models of animal contests such as the Hawk-Dove game predict that variation in fighting behavior will persist due to mixed evolutionarily stable strategies (ESS) under certain conditions. However, the genetic basis for this variation is poorly understood and a mixed ESS for fighting can be interpreted in more than one way. Specifically, we do not know whether variation in aggression within a population arises from among-individual differences in fixed strategy (determined by an individual's genotype—direct genetic effects [DGEs]), or from within-individual variation in strategy across contests. Furthermore, as suggested by developments of the original Hawk-Dove model, within-individual variation in strategy may be dependent on the phenotype and thus genotype of the opponent (indirect genetic effects—IGEs). Here we test for the effect of DGEs and IGEs during fights in the beadlet sea anemone Actinia equina. By exploiting the unusual reproductive system of sea anemones, combined with new molecular data, we investigate the role of both additive (DGE + IGE) and non-additive (DGE × IGE) genetic effects on fighting parameters, the latter of which have been hypothesized but never tested for explicitly. We find evidence for heritable variation in fighting ability and that fight duration increases with relatedness. Fighting success is influenced additively by DGEs and IGEs but we found no evidence for non-additive IGEs. These results indicate that variation in fighting behavior is driven by additive indirect genetic effects (DGE + IGE), and support a core assumption of contest theory that strategies are fixed by DGEs. [ABSTRACT FROM AUTHOR]

A key element at the heart of the Fisher–Lande–Kirkpatrick model of the sexual selection process is the genetic correlation between (male) trait and (female) preference. The strength of this association is critical in determining a population's evolutionary trajectory, which is why estimating its magnitude is so important. In the Lande model, the trait-preference correlation is solely established and maintained by mate choice, and although it is unclear how exclusively mate choice does this in nature, the experimental designs typically employed to measure trait-preference genetic correlations could be systematically weakening estimates by not allowing free mate choice (similarly with husbandry practices). The precise impact of the problem is unknown, and possibly unknowable, but simple solutions can be applied to ensure the accuracy of trait-preference correlation estimates is maximized. [ABSTRACT FROM AUTHOR]

Personality variation, defined as among‐individual differences in behaviour that are repeatable across time and context, is widely reported across animal taxa. From an evolutionary perspective, characterising the amount and structure of this variation is useful since differences among individuals are the raw material for adaptive behavioural evolution. However, behavioural variation among individuals also has implications for more applied areas of evolution and ecology—from invasion biology to ecotoxicology and selective breeding in captive systems. Here, we investigate the structure of personality variation in the red cherry shrimp, Neocaridina heteropoda, a popular ornamental species that is readily kept and bred under laboratory conditions and is emerging as a decapod crustacean model across these fields, but for which basic biological, ecological and behavioural data are limited. Using two assays and a repeated measures approach, we quantify behaviours putatively indicative of shy–bold variation and test for sexual dimorphism and/or size‐dependent behaviours (as predicted by some state‐dependent models of personality). We find moderate‐to‐high behavioural repeatabilities in most traits. Although strong individual‐level correlations across behaviours are consistent with a major personality axis underlying these observed traits, the multivariate structure of personality variation does not fully match a priori expectations of a shy–bold axis. This may reflect our ecological naivety with respect to what really constitutes bolder, more risk‐prone, behaviour in this species. We find no evidence for sexual dimorphism and only weak support for size‐dependent behaviour. Our study contributes to the growing literature describing behavioural variation in aquatic invertebrates. Furthermore, it lays a foundation for further studies harnessing the potential of this emerging model system. In particular, this existing behavioural variation could be functionally linked to life‐history traits and invasive success and serve as a target of artificial selection or bioassays. It thus holds significant promise in applied research across ecotoxicology, aquaculture and invasion biology. [ABSTRACT FROM AUTHOR]

Organisms can affect one another's phenotypes when they socially interact. Indirect genetic effects occur when an individual's phenotype is affected by genes expressed in another individual. These heritable effects can enhance or reduce adaptive potential, thereby accelerating or reversing evolutionary change. Quantifying these social effects is therefore crucial for our understanding of evolution, yet estimates of indirect genetic effects in wild animals are limited to dyadic interactions. We estimated indirect phenotypic and genetic effects, and their covariance with direct effects, for the date of spring breeding in North American red squirrels (Tamiasciurus hudsonicus) living in an array of territories of varying spatial proximity. Additionally, we estimated indirect effects and the strength of selection at low and high population densities. Social effects of neighbours on the date of spring breeding were different from zero at high population densities but not at low population densities. Indirect phenotypic effects accounted for a larger amount of variation in the date of breeding than differences attributable to the among‐individual variance, suggesting social interactions are important for determining breeding dates. The genetic component to these indirect effects was however not statistically significant. We therefore showcase a powerful and flexible method that will allow researchers working in organisms with a range of social systems to estimate indirect phenotypic and genetic effects, and demonstrate the degree to which social interactions can influence phenotypes, even in a solitary species. [ABSTRACT FROM AUTHOR]

Social interactions can give rise to indirect genetic effects (IGEs), which occur when genes expressed in one individual affect the phenotype of another individual. The evolutionary dynamics of traits can be altered when there are IGEs. Sex often involves indirect effects arising from first‐order (current) or second‐order (prior) social interactions, yet IGEs are infrequently quantified for reproductive behaviors. Here, we use experimental populations of burying beetles that have experienced bidirectional selection on mating rate to test for social plasticity and IGEs associated with focal males mating with a female either without (first‐order effect) or with (second‐order effect) prior exposure to a competitor, and resource defense behavior (first‐order effect). Additive IGEs were detected for mating rate arising from (first‐order) interactions with females. For resource defense behavior, a standard variance partitioning analysis provided no evidence of additive genetic variance—either direct or indirect. However, behavior was predicted by focal size relative to that of the competitor, and size is also heritable. Assuming that behavior is causally dependent on relative size, this implies that both DGEs and IGEs do occur (and may potentially interact). The relative contribution of IGEs may differ among social behaviors related to mating which has consequences for the evolutionary trajectories of multivariate traits. Behavioral traits involved in sexual selection can have similar phenotypically plastic responses to a change in social environment but may have quite different underlying architecture so will respond differently to selection. This highlights the importance of quantifying indirect genetic effects (IGEs) as well as direct genetic effects (DGEs) of suites of related traits to understand how they will evolve. [ABSTRACT FROM AUTHOR]

There is a systematic and suggestive analogy between grounding and causation. In my view, this analogy is no coincidence. Grounding and causation are alike because grounding is a type of causation: metaphysical causation. In this paper I defend the identification of grounding with metaphysical causation, drawing on the causation literature to explore systematic connections between grounding and metaphysical dependence counterfactuals, and I outline a non‐reductive counterfactual theory of grounding along interventionist lines. [ABSTRACT FROM AUTHOR]

This paper outlines a non‐reductive counterfactual account of grounding along interventionist lines, and uses the account to argue that taking grounding seriously requires ascribing non‐trivial truth‐conditions to a range of counterpossible counterfactuals. This result allows for a diagnosis of a route to scepticism about grounding, as deriving at least in part from scepticism about non‐trivial counterpossible truth and falsity. [ABSTRACT FROM AUTHOR]

Individual repeatability (R), defined as the proportion of observed variance attributable to among‐individual differences, is a widely used summary statistic in evolutionarily motivated studies of morphology, life history, physiology and, especially, behaviour. Although statistical methods to estimate R are well known and widely available, there is a growing tendency for researchers to interpret R in ways that are subtly, but importantly, different. Some view R as a property of a dataset and a statistic to be interpreted agnostically with respect to mechanism. Others wish to isolate the contributions of 'intrinsic' and/or 'permanent' individual differences, and draw a distinction between true (intrinsic) and pseudo‐repeatability arising from uncontrolled extrinsic effects. This latter view proposes a narrower, more mechanistic interpretation, than the traditional concept of repeatability, but perhaps one that allows stronger evolutionary inference as a consequence (provided analytical pitfalls are successfully avoided). Neither perspective is incorrect, but if we are to avoid confusion and fruitless debate, there is a need for researchers to recognise this dichotomy, and to ensure clarity in relation to how, and why, a particular estimate of R is appropriate in any case. [ABSTRACT FROM AUTHOR]

Analysis of data on school leavers in the UK points to considerable inequality in access to higher education. This is particularly acute in Scotland where access to the most competitive courses remains extremely difficult for those young people who are first in their generation to try and enter university or from low-income households. Programs to address this issue have had varied approaches framed around policy ambitions to “close the attainment gap” or “raise aspirations.” However, research has argued that these approaches have largely conceptualized the problem at an individual rather than structural level (Cummings et al., 2012; Gorard, Beng, & Davies, 2012). The Intergenerational Mentoring Network is an ongoing research and development project which recruits retired professionals as volunteer mentors and matches them with 15–18 year old secondary school pupils from lower-income areas of the city. This paper draws from research within this project to demonstrate the difficulties these young people face surrounding their interaction with very different social and cultural worlds, of which they have limited understanding and very restricted access. The paper illuminates the crucial role that intergenerational relationships can play in supporting young people in their journey into higher education. These are roles that, collectively, are enabling more young people to move into university and professional careers and through increasing social mobility are challenging inequality. [ABSTRACT FROM PUBLISHER]

The pace-of-life syndrome (POLS) hypothesis predicts associations between life history and 'risky' behaviours. Individuals with 'fast' lifestyles should develop faster, reproduce earlier, exhibit more risk-prone behaviours, and die sooner than those with 'slow' lifestyles. While support for POLS has been equivocal to date, studies have relied on individual-level (phenotypic) patterns in which genetic trade-offs may be masked by environmental effects on phenotypes. We estimated genetic correlations between life history (development, lifespan, size) and risky behaviours (exploration, aggression) in a pedigreed population of Mediterranean field crickets (Gryllus bimaculatus). Path analyses showed that behaviours mediated some genetic relationships between life history traits, though not those involved in trade-offs. Thus, while specific predictions of POLS theory were not supported, genetic integration of behaviour and life history was present. This implies a major role for risky behaviours in life history evolution. [ABSTRACT FROM AUTHOR]

Inequality in educational attainment is a serious and increasing problem in Scotland and the rest of the United Kingdom, and youth mentoring is gaining popularity as an intervention to address this issue. However, evidence for the effectiveness of mentoring is weak, and research suggests that a large proportion of mentoring relationships end prematurely, which can be harmful to young people. This article draws on data from the Glasgow Intergenerational Mentoring Network, an ongoing research and development project facilitating mentoring for secondary school pupils in socially disadvantaged areas, employing retired professionals as volunteer mentors. Case material is presented to discuss the nature of mentoring and the role of the mentor. This article conceptualizes volunteer mentors as relational experts, relational learners, and nonrelational mentors. The implications of these categories are discussed in terms of recruitment, training, and support of mentors, to facilitate ethical and effective mentoring relationships. [ABSTRACT FROM AUTHOR]

Having recognized that variation around the population-level "Golden Mean" of labile traits contains biologically meaningful information, behavioural ecologists have focused increasingly on exploring the causes and consequences of individual variation in behaviour. These are exciting new directions for the field, assisted in no small part by the adoption of mixed-effects modelling techniques that enable the partitioning of among- and within-individual behavioural variation. It has become commonplace to extract predictions of individual random effects from such models for use in subsequent analyses (for example, between a personality trait and other individual traits such as cognition, physiology, or fitness-related measures). However, these predictions are made with large amounts of error that is not carried forward, rendering further tests susceptible to spurious P values from these individual-level point estimates. We briefly summarize the problems with such statistical methods that are used regularly by behavioural ecologists, and highlight the robust solutions that exist within the mixed model framework, providing tutorials to aid in their implementation. [ABSTRACT FROM AUTHOR]

If the most familiar overlapping (branching universe) interpretation of Everettian quantum mechanics (EQM) is correct, then each of us is constantly splitting into multiple people. This consequence gives rise to thequantum doomsday argument, which threatens to draw crippling epistemic consequences from EQM. However, a diverging (parallel universe) interpretation of EQM undermines the quantum doomsday argument completely. This appears to tell in favour of the diverging interpretation. But it is surprising that a metaphysical question that is apparently underdetermined by the physics should be settled by purely epistemological considerations; and I argue that the positive case for divergence based on the quantum doomsday effect is ultimately unsuccessful. I discuss how some influential treatments of Everettian confirmation handle the quantum doomsday puzzle, and suggest that it can most satisfyingly be resolved via a naturalistic approach to the metaphysics of modality. [ABSTRACT FROM AUTHOR]

Some species actively change color and pattern for camouflage on a range of background types. Such dynamic camouflage may be particularly advantageous for species inhabiting heterogeneous habitats, such as intertidal zones, where individuals are exposed to both terrestrial and marine predators depending on tides and wave action. Most studies of dynamic pattern camouflage have focused on relatively few species, and rarely species inhabiting the intertidal zone. We used image analysis and predator (avian) vision modelling to determine if rock gobies (Gobius paganellus) change their body pattern in response to their background, and to explore how background marking size influence pattern change. Rock gobies rapidly (within 1 min) changed their pattern when placed on checkerboards with different sized squares, and on backgrounds resembling natural substrates. On backgrounds resembling natural substrates, those with a small grain size, such as sand, elicited a larger degree of pattern change than those with a larger grain size. However, despite this, the majority of fish showed little or no improvement in background matching over time. Instead, the markings elicited are characteristic of disruptive coloration and may function primarily through breaking up the body outline rather than via improved match to the background pattern itself. [ABSTRACT FROM AUTHOR]

In animal contests, individuals respond plastically to the phenotypes of the opponents that they confront. These "opponent"—or "indirect"—effects are often repeatable, for example, certain opponents consistently elicit more or less aggressiveness in others. "Personality" (repeatable among-individual variation in behavior) has been proposed as an important source of indirect effects. Here, we repeatedly assayed aggressiveness of wild-caught adult male field crickets Gryllus campestris in staged dyadic fights, measuring aggressiveness of both contestants. Measurements of their personality in nonsocial contexts (activity and exploration behavior) enabled us to ask whether personality caused indirect effects on aggressiveness. Activity, exploration, and aggressiveness were positively associated into a behavioral syndrome eliciting aggressiveness in conspecifics, providing direct evidence for the role of personality in causing indirect effects. Our findings imply that a multivariate view of phenotypes that includes indirect effects greatly improves our ability to understand the ecology and evolution of behavior. [ABSTRACT FROM AUTHOR]

Innovative behaviour may allow animals to cope with changes in their environment. Innovative propensities are known to vary widely both between and within species, and a growing body of research has begun to examine the factors that drive individuals to innovate. Evidence suggests that individuals are commonly driven to innovate by necessity; for instance by hunger or because they are physically unable to outcompete others for access to resources. However, it is not known whether the factors that drive individuals to innovate are stable across contexts. We examined contextual variation in the drivers of innovation in rock pool prawns (Palaemon spp), invertebrates that face widely fluctuating environments and may, through the actions of tides and waves, find themselves isolated or in groups. Using two novel foraging tasks, we examined the effects of body size and hunger in prawns tested in solitary and group contexts. When tested alone, small prawns were significantly more likely to succeed in a spatial task, and faster to reach the food in a manipulation task, while hunger state had no effect. In contrast, size had no effect when prawns were tested in groups, but food-deprived individuals were disproportionately likely to innovate in both tasks. We suggest that contextual variation in the drivers of innovation is likely to be common in animals living in variable environments, and may best be understood by considering variation in the perception of relative risks and rewards under different conditions. [ABSTRACT FROM AUTHOR]

Background: Life-history studies of wild bird populations often focus on the relationship between an individual’s condition and its capacity to mount an immune response, as measured by a commonly-employed assay of cutaneous immunity, the PHA skin test. In addition, haematocrit, the packed cell volume in relation to total blood volume, is often measured as an indicator of physiological performance. A multi-year study of a wild population of house wrens has recently revealed that those exhibiting the highest condition and strongest PHA responses as nestlings are most likely to be recruited to the breeding population and to breed through two years of age; in contrast, intermediate haematocrit values result in the highest recruitment to the population. Selection theory would predict, therefore, that most of the underlying genetic variation in these traits should be exhausted resulting in low heritability, although such traits may also exhibit low heritability because of increased residual variance. Here, we examine the genetic and environmental variation in condition, cutaneous immunity, and haematocrit using an animal model based on a pedigree of approximately 2,800 house wrens. Results: Environmental effects played a paramount role in shaping the expression of the fitness-related traits measured in this wild population, but two of them, condition and haematocrit, retained significant heritable variation. Condition was also positively correlated with both the PHA response and haematocrit, but in the absence of any significant genetic correlations, it appears that this covariance arises through parallel effects of the environment acting on this suite of traits. Conclusions: The maintenance of genetic variation in different measures of condition appears to be a pervasive feature of wild bird populations, in contradiction of conventional selection theory. A major challenge in future studies will be to explain how such variation persists in the face of the directional selection acting on condition in house wrens and other species. [ABSTRACT FROM AUTHOR]

Darren Bradley has recently appealed to observation selection effects to argue that conditionalization presents no special problem for Everettian quantum mechanics, and to defend the ‘halfer’ answer to the puzzle of Sleeping Beauty. I assess Bradley’s arguments and conclude that while he is right about confirmation in Everettian quantum mechanics, he is wrong about Sleeping Beauty. This result is doubly good news for Everettians: they can endorse Bayesian confirmation theory without qualification, but they are not thereby compelled to adopt the unpopular ‘halfer’ answer in Sleeping Beauty. These considerations suggest that objective chance is playing an important and under-appreciated role in Sleeping Beauty.1 Introduction2 Confirmation in Everettian Quantum Mechanics3 Sleeping Beauty4 The Selection Model5 Bradley’s Argument6 The Right Route to ⅓7 The Breakdown of the Analogy8 Alternative Diagnoses9 God’s Gambling Game10 Non-chancy Sleeping Beauty Cases11 Conclusion [ABSTRACT FROM AUTHOR]

David Wallace has given a decision-theoretic argument for the Born Rule in the context of Everettian quantum mechanics (EQM). This approach promises to resolve some long-standing problems with probability in EQM, but it has faced plenty of resistance. One kind of objection (the ‘incoherence problem’) charges that the requisite notion of decision-theoretic uncertainty is unavailable in the Everettian picture, so that the argument cannot gain any traction; another kind of objection grants the proof’s applicability and targets the premises. In this article I propose some novel principles connecting the physics of EQM with the metaphysics of modality, and argue that in the resulting framework the incoherence problem does not arise. These principles also help to justify one of the most controversial premises of Wallace’s argument, ‘branching indifference’. Absent any a priori reason to align the metaphysics with the physics in some other way, the proposed principles can be adopted on grounds of theoretical utility. The upshot is that Everettians can, after all, make clear sense of objective probability. 1 Introduction2 Setup3 Individualism versus Collectivism4 The Ingredients of Indexicalism5 Indexicalism and Incoherence 5.1 The trivialization problem 5.2 The uncertainty problem6 Indexicalism and Branching Indifference 6.1 Introducing branching indifference 6.2 The pragmatic defence of branching indifference 6.3 The non-existence defence of branching indifference 6.4 The indexicalist defence of branching indifference7 Conclusion [ABSTRACT FROM PUBLISHER]

In 'Quiddistic Knowledge' (Schaffer in Philos Stud 123:1-32, ), Jonathan Schaffer argued influentially against the view that the laws of nature are metaphysically necessary. In this reply I aim to show how a coherent and well-motivated form of necessitarianism can withstand his critique. Modal necessitarianism-the view that the actual laws are the laws of all possible worlds-can do justice to some intuitive motivations for necessitarianism, and it has the resources to respond to all of Schaffer's objections. It also has certain advantages over contingentism in the domain of modal epistemology. I conclude that necessitarianism about laws remains a live option. [ABSTRACT FROM AUTHOR]

Although senescence in reproductive success has been observed in a number of wild animal populations, longitudinal analyses determining the actual processes responsible for population-level patterns of ageing, and in particular, the relative contributions of within-individual senescence vs. selective mortality, are rare. Furthermore, many studies also only consider single traits in isolation, despite evidence that different aspects of physiology and fitness may show varying patterns of senescence., We used data from an unmanaged population of Soay sheep on the islands of St Kilda, NW Scotland to analyse age-related change in four aspects of female reproductive performance: annual fecundity, twinning rate and the maternally influenced offspring traits of lamb birth weight and early survival. We present the results of three analytical techniques, which attempt to disentangle the contributions of within-individual ageing, selective mortality and terminal effects to the observed population-level patterns of age-specific change., All four traits showed considerable age-related change, but population-level trajectories varied markedly across them. Population-level patterns were underpinned by considerable within-individual senescence including within-individual terminal declines in performance occurring only in the final year of life. However, at the population level, these processes were also masked to some extent by selective disappearance (mortality) of poor breeders., The contributions of individual-level ageing and selective mortality to population-level ageing patterns varied considerably between traits, as did the nature of individual-level senescence. We did not observe detectable significant late-life declines in all four traits: in particular, there was no evidence of senescence in twinning rate. The results indicate that senescence may not be ubiquitous across all aspects of reproductive performance within a population. [ABSTRACT FROM AUTHOR]

The guppy Y chromosome has been considered a model system for the evolution of suppressed recombination between sex chromosomes, and it has been proposed that complete sex‐linkage has evolved across about 3 Mb surrounding this fish's sex‐determining locus, followed by recombination suppression across a further 7 Mb of the 23 Mb XY pair, forming younger "evolutionary strata". Sequences of the guppy genome show that Y is very similar to the X chromosome. Knowing which parts of the Y are completely nonrecombining, and whether there is indeed a large completely nonrecombining region, are important for understanding its evolution. Here, we describe analyses of PoolSeq data in samples from within multiple natural populations from Trinidad, yielding new results that support previous evidence for occasional recombination between the guppy Y and X. We detected recent demographic changes, notably that downstream populations have higher synonymous site diversity than upstream ones and other expected signals of bottlenecks. We detected evidence of associations between sequence variants and the sex‐determining locus, rather than divergence under a complete lack of recombination. Although recombination is infrequent, it is frequent enough that associations with SNPs can suggest the region in which the sex‐determining locus must be located. Diversity is elevated across a physically large region of the sex chromosome, conforming to predictions for a genome region with infrequent recombination that carries one or more sexually antagonistic polymorphisms. However, no consistently male‐specific variants were found, supporting the suggestion that any completely sex‐linked region may be very small. [ABSTRACT FROM AUTHOR]

Quantitative genetic studies in natural populations are of growing interest to speciation research since divergence is often believed to arise through micro-evolutionary change, caused by natural selection on functional morphological traits. The species flock of cichlid fishes in Africa's oldest lake, Lake Tanganyika, offers a rare opportunity to study this process. Using the cichlid species Tropheus moorii, we assessed the potential for microevolution in a set of morphological traits by estimating their quantitative genetic basis of variation. Two approaches were employed: (1) estimation of trait heritabilities ( h) in situ from a sample of wild caught fish, and (2) estimation of h from first generation offspring produced in a semi-natural breeding experiment. In both cases, microsatellite data were used to infer pedigree structure among the sampled individuals and estimates of h were made using an animal model approach. Although power was limited by the pedigree structures estimated (particularly in the wild caught sample), we nonetheless demonstrate the presence of significant additive genetic variance for aspects of morphology that, in the cichlid species Tropheus moorii, are expected to be functionally and ecologically important, and therefore likely targets of natural selection. We hypothesize that traits showing significant additive genetic variance, such as the mouth position have most likely played a key role in the adaptive evolution of the cichlid fish Tropheus moorii. [ABSTRACT FROM AUTHOR]

Aggression occurs when individuals compete over limiting resources. While theoretical studies have long placed a strong emphasis on context-specificity of aggression, there is increasing recognition that consistent behavioural differences exist among individuals, and that aggressiveness may be an important component of individual personality. Though empirical studies tend to focus on one aspect or the other, we suggest there is merit in modelling both within- and among-individual variation in agonistic behaviour simultaneously. Here, we demonstrate how this can be achieved using multivariate linear mixed effect models. Using data from repeated mirror trials and dyadic interactions of male green swordtails, Xiphophorus helleri, we show repeatable components of (co)variation in a suite of agonistic behaviour that is broadly consistent with a major axis of variation in aggressiveness. We also show that observed focal behaviour is dependent on opponent effects, which can themselves be repeatable but were more generally found to be context specific. In particular, our models show that within-individual variation in agonistic behaviour is explained, at least in part, by the relative size of a live opponent as predicted by contest theory. Finally, we suggest several additional applications of the multivariate models demonstrated here. These include testing the recently queried functional equivalence of alternative experimental approaches, (e.g., mirror trials, dyadic interaction tests) for assaying individual aggressiveness. [ABSTRACT FROM AUTHOR]

Simon Saunders and David Wallace have proposed an attractive semantics for interpreting linguistic communities embedded in an Everettian multiverse. It provides a charitable interpretation of our ordinary talk about the future, and allows us to retain a principle of bivalence for propositions and to retain the law of excluded middle in the logic of propositions about the future. But difficulties arise when it comes to providing an appropriate account of the metaphysics of macroscopic objects and events. I evaluate various metaphysical frameworks which might be combined with the Saunders-Wallace semantics. I conclude that the most appropriate metaphysics to underwrite the semantics renders Everettian quantum mechanics a theory of non-overlapping worlds. [ABSTRACT FROM AUTHOR]

Cryptic evolution has been defined as adaptive evolutionary change being masked by concurrent environmental change. Empirical studies of cryptic evolution have usually invoked a changing climate and/ or increasing population density as the form of detrimental environmental change experienced by a population undergoing cryptic evolution. However, Fisher (1958) emphasized that evolutionary change in itself is likely to be an important component of "environmental deterioration," a point restated by Cooke et al. (1990) in the context of intraspecific competition. In this form, environmental deterioration arises because a winning lineage has to compete against more winners in successive generations as the population evolves. This "evolutionary environmental deterioration" has different implications for the selection and evolution of traits influenced by resource competition than general environmental change. We reformulate Cooke's model as a quantitative genetic model to show that it is identical in form to more recent developments proposed by quantitative geneticists. This provides a statistical framework for discriminating between the alternative hypotheses of environmental change and environmental deterioration caused by evolutionary change. We also demonstrate that in systems where no phenotypic change has occurred, there are many reasonable biological processes that will generate patterns in predicted breeding values that are consistent with what has been interpreted as cryptic evolution, and care needs to be taken when interpreting these patterns. These processes include mutation, sib competition, and invisible fractions. [ABSTRACT FROM AUTHOR]

Victims of infection are expected to suffer increasingly as parasite population growth increases. Yet, under some conditions, faster-growing parasites do not appear to cause more damage, and infections can be quite tolerable. We studied these conditions by assessing how the relationship between parasite population growth and host health is sensitive to environmental variation. In experimental infections of the crustacean Daphnia magna and its bacterial parasite Pasteuria ramosa, we show how easily an interaction can shift from a severe interaction, that is, when host fitness declines substantially with each unit of parasite growth, to a tolerable relationship by changing only simple environmental variables: temperature and food availability. We explored the evolutionary and epidemiological implications of such a shift by modeling pathogen evolution and disease spread under different levels of infection severity and found that environmental shifts that promote tolerance ultimately result in populations harboring more parasitized individuals. We also find that the opportunity for selection, as indicated by the variance around traits, varied considerably with the environmental treatment. Thus, our results suggest two mechanisms that could underlie coevolutionary hotspots and coldspots: spatial variation in tolerance and spatial variation in the opportunity for selection. [ABSTRACT FROM AUTHOR]

There is now overwhelming evidence concerning the awareness people with moderate to mild intellectual disabilities have about the stigma they experience in their lives. However, there is still controversy about the potential impact of stigma on their self-perceptions. This article will draw on findings from an ethnographic study to show that even when individuals have difficulty expressing their views verbally, their actions can provide evidence of how they struggle to establish or maintain positive social identities—sometimes at the cost of their mental health. The implications of these and other findings will be discussed in relation to social constructionist theories of self-perception. This in turn will be linked to a discussion about the kind of support that might be required by people with intellectual disabilities, and how stigma might increase vulnerability to emotional and interpersonal problems. [ABSTRACT FROM AUTHOR]

How should we respond to cases of disagreement where two epistemic agents have the same evidence but come to different conclusions? Adam Elga has provided a Bayesian framework for addressing this question. In this paper, I shall highlight two unfortunate consequences of this framework, which Elga does not anticipate. Both problems derive from a failure of commutativity between application of the equal weight view and updating in the light of other evidence. [ABSTRACT FROM AUTHOR]

Analyses of pedigrees and pedigree-derived parameters (e.g. relatedness and fitness) provide some of the most informative types of studies in evolutionary biology. Ther packagepedantics implements tools to facilitate power and sensitivity analyses of pedigree-related studies of natural populations. Functions are available to permute pedigree data in various ways with the goal of mimicking patterns of pedigree errors and missingness that occur in studies of natural populations. Another set of functions simulates genetic and phenotypic data based on arbitrary pedigrees. Finally, functions are also available with which visual and numerical representations of pedigree structure can be generated. [ABSTRACT FROM AUTHOR]

In studies of population ecology, demography and life history evolution, among-individual differences in traits associated with survival and reproduction are often attributed to variation in ‘individual quality’. However, often intuitive quality is rarely defined explicitly, and we argue that this can result in ambiguity about what quality actually is. Here we consider the various ways in which the concept of quality is currently applied, and show that subtle differences in intended meaning have very important consequences when the goal is to draw evolutionary inferences. We also propose a novel approach that is consistent with all current ecological uses, but also allows the concept of quality to be integrated with existing evolutionary theory. [Copyright &y& Elsevier]

The independent evolution of the sexes may often be constrained if male and female homologous traits share a similar genetic architecture. Thus, cross-sex genetic covariance is assumed to play a key role in the evolution of sexual dimorphism (SD) with consequent impacts on sexual selection, population dynamics, and speciation processes. We compiled cross-sex genetic correlations ( rMF) estimates from 114 sources to assess the extent to which the evolution of SD is typically constrained and test several specific hypotheses. First, we tested if rMF differed among trait types and especially between fitness components and other traits. We also tested the theoretical prediction of a negative relationship between rMF and SD based on the expectation that increases in SD should be facilitated by sex-specific genetic variance. We show that rMF is usually large and positive but that it is typically smaller for fitness components. This demonstrates that the evolution of SD is typically genetically constrained and that sex-specific selection coefficients may often be opposite in sign due to sub-optimal levels of SD. Most importantly, we confirm that sex-specific genetic variance is an important contributor to the evolution of SD by validating the prediction of a negative correlation between rMF and SD. [ABSTRACT FROM AUTHOR]

1. Efforts to understand the links between evolutionary and ecological dynamics hinge on our ability to measure and understand how genes influence phenotypes, fitness and population dynamics. Quantitative genetics provides a range of theoretical and empirical tools with which to achieve this when the relatedness between individuals within a population is known. 2. A number of recent studies have used a type of mixed-effects model, known as the animal model, to estimate the genetic component of phenotypic variation using data collected in the field. Here, we provide a practical guide for ecologists interested in exploring the potential to apply this quantitative genetic method in their research. 3. We begin by outlining, in simple terms, key concepts in quantitative genetics and how an animal model estimates relevant quantitative genetic parameters, such as heritabilities or genetic correlations. 4. We then provide three detailed example tutorials, for implementation in a variety of software packages, for some basic applications of the animal model. We discuss several important statistical issues relating to best practice when fitting different kinds of mixed models. 5. We conclude by briefly summarizing more complex applications of the animal model, and by highlighting key pitfalls and dangers for the researcher wanting to begin using quantitative genetic tools to address ecological and evolutionary questions. [ABSTRACT FROM AUTHOR]

Best linear unbiased prediction (BLUP) is a method for obtaining point estimates of a random effect in a mixed effect model. Over the past decade it has been used extensively in ecology and evolutionary biology to predict individual breeding values and reaction norms. These predictions have been used to infer natural selection, evolutionary change, spatial-genetic patterns, individual reaction norms, and frailties. In this article we show analytically and through simulation and example why BLUP often gives anticonservative and biased estimates of evolutionary and ecological parameters. Although some concerns with BLUP methodology have been voiced before, the scale and breadth of the problems have probably not been widely appreciated. Bias arises because BLUPs are often used to estimate effects that are not explicitly accounted for in the model used to make the predictions. In these cases, predicted breeding values will often say more about phenotypic patterns than the genetic patterns of interest. An additional problem is that BLUPs are point estimates of quantities that are usually known with little certainty. Failure to account for this uncertainty in subsequent tests can lead to both bias and extreme anticonservatism.We demonstrate that restricted maximum likelihood and Bayesian solutions exist for these problems and show how unbiased and powerful tests can be derived that adequately quantify uncertainty. Of particular utility is a new test for detecting evolutionary change that not only accounts for prediction error in breeding values but also accounts for drift. To illustrate the problem, we apply these tests to long-term data on the Soay sheep (Ovis aries) and the great tit (Parus major) and show that previously reported temporal trends in breeding values are not supported. [ABSTRACT FROM AUTHOR]

This work demonstrates that environmental conditions experienced by individuals can shape their development and affect the stability of genetic associations. The implication of this observation is that the environmental response may influence the evolution of traits in the wild. Here, we examined how the genetic architecture of a suite of sexually dimorphic traits changed as a function of environmental conditions in an unmanaged population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, northwest Scotland. We examined the stability of phenotypic, genetic, and environmental (residual) covariance in males during the first year of life between horn length, body weight, and parasite load in environments of different quality. We then examined the same covariance structures across environments within and between the adult sexes. We found significant genotype-by-environment interactions for lamb male body weight and parasite load, leading to a change in the genetic correlation among environments. Horn length was genetically correlated with body weight in males but not females and the genetic correlation among traits within and between the sexes was dependent upon the environmental conditions experienced during adulthood. Genetic correlations were smaller in more favorable environmental conditions, suggesting that in good environ- merits, loci are expressed that have sex-specific effects. The reduction in genetic correlation between the sexes may allow independent evolutionary trajectories for each sex. This study demonstrates that the genetic architecture of traits is not stable under temporally varying environments and highlights the fact that evolutionary processes may depend largely upon ecological conditions. [ABSTRACT FROM AUTHOR]

FST and RST estimates for Arctic charr from six microsatelite markers collected from two neighbouring Scottish lakes, Loch Maree and Loch Stack, confirm the presence of two distinct genetic groupings representing separate populations within each lake. In both lakes, there was also a clear body size dimorphism, with large and small body size forms that segregated according to genetic grouping. There was evidence of only subtle foraging ecology differences between morphs, with the small body size morph in both lakes being more generalist in its foraging in the summer (consuming mostly plankton but also some macrobenthos) than the large body size morph, which specialized on planktonic prey. Trophic morphology (head and mouth shape) did not differ significantly between morphs (although the small sample size for Maree makes this a preliminary finding). Cluster analysis of the microsatelite data and the presence of private alleles showed that morphologically similar forms in different lakes were not genetically similar, as would be expected under a multiple invasion hypothesis. Thus, the data do not support a hypothesis of a dual invasion of both lakes by two common ancestors but instead suggest an independent origin of the two forms in each lake. Thus parallel sympatric divergence as a result of common selection pressures in both lakes is the most parsimonious explanation of the evolutionary origin of these polymorphisms. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 748–757. [ABSTRACT FROM AUTHOR]

Summary: Aging, or senescence, defined as a decline in physiological function with age, has long been a focus of research interest for evolutionary biologists. How has natural selection failed to remove genetic effects responsible for such reduced fitness among older individuals? Current evolutionary theory explains this phenomenon by showing that, as a result of the risk of death from environmental causes that individuals experience, the force of selection inevitably weakens with age . This in turn means that genetic mutations having detrimental effects that are only felt late in life might persist in a population. Although widely accepted, this theory rests on the assumption that there is genetic variation for aging in natural systems , or (equivalently), that genotype-by-age interactions (GxA) occur for fitness. To date, empirical support for this assumption has come almost entirely from laboratory studies on invertebrate systems, most notably Drosophila and C. elegans, whereas tests of genetic variation for aging are largely lacking from natural populations . By using data from two wild mammal populations, we perform quantitative genetic analyses of fitness and provide the first evidence for a genetic basis of senescence to come from a study in the natural environment. We find evidence that genetic differences among individuals cause variation in their rates of aging and that additive genetic variance for fitness increases with age, as predicted by the evolutionary theory of senescence. [Copyright &y& Elsevier]

Senescence is the decline in survival and reproduction as an organism ages and is known to occur in collared flycatchers Ficedula albicollis. We consider annual fitness (the estimated genetic contribution that an individual makes to next year's gene pool) as a measure of age-specific fitness. We apply a restricted maximum likelihood linear mixed-model approach on 25 years of data on 3,844 male and 4,992 female collared flycatchers. Annual fitness had a significant additive genetic component (h2 of about 4%). Annual fitness declined at later ages in both sexes. Using a random regression animal model, we show that the observed age-related phenotypic changes in annual fitness were not present on the additive genetic level, contrary to predictions of genetic hypotheses of senescence. Our study suggests that patterns of aging in the wild need to be interpreted with caution in terms of underlying genetics because they may be largely determined by environmental processes. [ABSTRACT FROM AUTHOR]

Evolution of size and growth depends on heritable variation arising from additive and maternal genetic effects. Levels of heritable (and nonheritable) variation might change over ontogeny, increasing through "variance compounding" or decreasing through "compensatory growth." We test for these processes using a meta-analysis of age-specific weight traits in domestic ungulates. Generally, mean standardized variance components decrease with age, consistent with compensatory growth. Phenotypic convergence among adult sheep occurs through decreasing environmental and maternal genetic variation. Maternal variation similarly declines in cattle. Maternal genetic effects are thus reduced with age (both in absolute and relative terms). Significant trends in heritability (decreasing in cattle, increasing in sheep) result from declining maternal and environmental components rather than from changing additive variation. There was no evidence for increasing standardized variance components. Any compounding must therefore be masked by more important compensatory processes. While extrapolation of these patterns to processes in natural population is difficult, our results highlight the inadequacy of assuming constancy in genetic parameters over ontogeny. Negative covariance between direct and maternal genetic effects was common. Negative correlations with additive and maternal genetic variances indicate that antagonistic pleiotropy (between additive and maternal genetic effects) may maintain genetic variance and limit responses to selection. [ABSTRACT FROM AUTHOR]

Body size is an important determinant of fitness in many organisms. While size will typically change over the lifetime of an individual, heritable components of phenotypic variance may also show ontogenetic variation. We estimated genetic (additive and maternal) and environmental covariance structures for a size trait (June weight) measured over the first 5 years of life in a natural population of bighorn sheep Ovis canadensis. We also assessed the utility of random regression models for estimating these structures. Additive genetic variance was found for June weight, with heritability increasing over ontogeny because of declining environmental variance. This pattern, mirrored at the phenotypic level, likely reflects viability selection acting on early size traits. Maternal genetic effects were significant at ages 0 and 1, having important evolutionary implications for early weight, but declined with age being negligible by age 2. Strong positive genetic correlations between age-specific traits suggest that selection on June weight at any age will likely induce positively correlated responses across ontogeny. Random regression modeling yielded similar results to traditional methods. However, by facilitating more efficient data use where phenotypic sampling is incomplete, random regression should allow better estimation of genetic (co)variances for size and growth traits in natural populations. [ABSTRACT FROM AUTHOR]

Genotypic errors, whether due to mutation or laboratory error, can cause the genotypes of parents and their offspring to appear inconsistent with Mendelian inheritance. As a result, molecular parentage analyses are expected to benefit when allowances are made for the presence of genotypic errors. However, a cost of allowing for genotypic errors might also be expected under some analytical conditions, primarily because parentage analyses that assume nonzero genotypic error rates can neither assign nor exclude parentage with certainty. The goal of this work was therefore to determine whether or not such costs might be important under conditions relevant to parentage analyses, particularly in natural populations. Simulation results indicate that the costs may often outweigh the benefits of accounting for nonzero error rates, except in situations where data are available for many marker loci. Consequently, the most powerful approach to handling genotypic errors in parentage analyses might be to apply likelihood equations with error rates set to values substantially lower than the rates at which genotypic errors occur. When applying molecular parentage analyses to natural populations, we advocate an increased consideration of optimal strategies for handling genotypic errors. Currently available software packages contain procedures that can be used for this purpose. [ABSTRACT FROM AUTHOR]

Reproductive and early life-history traits can be considered aspects of either offspring or maternal phenotype, and their evolution will therefore depend on selection operating through offspring and maternal components of fitness. Furthermore, selection at these levels may be antagonistic, with optimal offspring and maternal fitness occurring at different phenotypic values. We examined selection regimes on the correlated traits of birth weight, birth date, and litter size in Soay sheep (Ovis aries) using data from a long-term study of a free-living population on the archipelago of St. Kilda, Scotland. We tested the hypothesis that selective constraints on the evolution of the multivariate phenotype arise through antagonistic selection, either acting at offspring and maternal levels, or on correlated aspects of phenotype. All three traits were found to be under selection through variance in short-term and lifetime measures of fitness. Analysis of lifetime fitness revealed strong positive directional selection on birth weight and weaker selection for increased birth date at both levels. However, there was also evidence for stabilizing selection on these traits at the maternal level, with reduced fitness at high phenotypic values indicating lower phenotypic optima for mothers than for offspring. Additionally, antagonistic selection was found on litter size. From the offspring's point of view it is better to be born a singleton, whereas maternal fitness increases with average litter size. The decreased fitness of twins is caused by their reduced birth weight; therefore, this antagonistic selection likely results from trade- offs between litter size and birth weight that have different optimal resolutions with respect to offspring and maternal fitness. Our results highlight how selection regimes may vary depending on the assignment of reproductive and early life-history traits to either offspring or maternal phenotype. [ABSTRACT FROM AUTHOR]

Supported employment is now regarded as a major vehicle for enabling people with learning difficulties to enter employment. The purpose of this paper is to use the evidence provided by three case studies of people with learning difficulties who have participated in supported employment to critically examine its fundamental premises. The paper argues that the influences of normalisation theory and the US model of supported employment have combined in the UK to form a variant of supported employment which, in pursuing the concept of real job, fails to adequately address the consequences of impairment. [ABSTRACT FROM AUTHOR]

The article presents information on implications of social capital theory for men and women with learning difficulties. It focuses on gender, social capital and lifelong learning for people with mental difficulties. Social capital is currently a very popular concept in social science. The social capital thinking is useful in understanding the nature and effects of people's social networks. The social capital offers a way of humanizing an essentially brutal economic system. The social capital focuses on the intervening variables of gender, social class and disability. Social capital within the family that gives the child access to the adult's human capital depends both on the physical presence of adults in the family and on the attention given by adults to the child. Single-parent families are observed to be deficient in social capital. The Learning Society project also presented the ways in which experiences of training and supported employment were structured by gender. The lack of access to coherent post-school education, training and employment, has negative consequences for the social capital of people with learning difficulties.

This article, drawing on the research findings from the ESRC-funded project 'The Meaning of the Learning Society for Adults with Learning Difficulties' explores how the introduction of the 1990 NHS and Community Care Act, with its emphasis on the establishment of a 'mixed economy' in social care, has affected the lives of people with learning difficulties living in a rural area of Scotland. The impact of the quasi-market, in particular the purchaser-provider split and the operation of contracts, on the lives of three adults with learning difficulties is examined. [ABSTRACT FROM AUTHOR]