Coefficients of relationship and coefficients of relatedness in kin selection: A covariance form for the RHO formula

Hamilton used the coefficient of relationship to predict the direction of selection acting on “genes for altruism” in a theoretical model. The sense (direction) of the inequality determined the direction of selection. But, paradoxically, Hamilton pointed out that traditional formulae for the coefficient of relationship cannot be evaluated during directional selection. Hamilton escaped the paradox by assuming very weak selection.Later papers, including three by Hamilton, used other coefficients in place of the coefficient of relationship in the inequality. Thus inbreeding and directional selection could be handled in exact models. This paper tries to clarify the differences between the various coefficients and to point out an error in one of the author's previous works: Namely to state that there is analytical proof that the regression coefficient is a special case of a coefficient called ? whereas the earlier paper claimed only numerical verification was possible.Since the symbol “?” is also used to denote the correlation coefficient, it is suggested here that the p of kin selection be given a new name.     

Social life: the paradox of multiple-queen colonies

The evolution of animal societies in which some individuals forego their own reproductive opportunities to help others to reproduce poses an evolutionary paradox that can be traced to Darwin. Altruism may evolve through kin selection when the donor and recipient of altruistic acts are related to each other, as generally is the case in social birds and mammals. Similarly, social insect workers are highly related to the brood they rear when colonies are headed by a single queen. However, recent studies have shown that insect colonies frequently contain several queens, with the effect of decreasing relatedness among colony members. How can one account for the origin and maintenance of such colonies? This evolutionary enigma presents many of the same theoretical challenges as does the evolution of cooperative breeding and eusociality.     

To perceive is to doubt: The relativity of perception

The uncertainty or entropy theory of perception is founded on the premise that for perception to occur, there must first of all be uncertainty. That is, perception or awareness is relative to the expectation of the perceiver. This view of perception leads to a seeming-paradox. How can there be uncertainty unless the alternatives have previously been perceived? But, by the premise of the theory, how can the alternatives have been perceived unless there was prior uncertainty? It is shown that this paradox may result physiologically in the concurrence of sensory and motor (or “active”) events during the process of perceiving. It is shown, further, that a close analogy exists between systems of formal logic and systems which perceive through uncertainty. This, in turn, suggests a basis for a calculus of perception.     

Constraints on the evolution of asexual reproduction

Sexual reproduction is almost ubiquitous among multicellular organisms even though it entails severe fitness costs. To resolve this apparent paradox, an extensive body of research has been devoted to identifying the selective advantages of recombination that counteract these costs. Yet, how easy is it to make the transition to asexual reproduction once sexual reproduction has been established for a long time? The present review approaches this question by considering factors that impede the evolution of parthenogenesis in animals. Most importantly, eggs need a diploid chromosome set in most species in order to develop normally. Next, eggs may need to be activated by sperm, and sperm may also contribute centrioles and other paternal factors to the zygote. Depending on how diploidy is achieved mechanistically, further problems may arise in offspring that stem from 'inbreeding depression' or inappropriate sex determination systems. Finally, genomic imprinting is another well-known barrier to the evolution of asexuality in mammals. Studies on species with occasional, deficient parthenogenesis indicate that the relative importance of these constraints may vary widely. The intimate evolutionary relations between haplodiploidy and parthenogenesis as well as implications for the clade selection hypothesis of the maintenance of sexual reproduction are also discussed.     

Is there a star tree paradox?

Concerns have been raised that posterior probabilities on phylogenetic trees can be unreliable when the true tree is unresolved or has very short internal branches, because existing methods for Bayesian phylogenetic analysis do not explicitly evaluate unresolved trees. Two recent papers have proposed that evaluating only resolved trees results in a "star tree paradox": when the true tree is unresolved or close to it, posterior probabilities were predicted to become increasingly unpredictable as sequence length grows, resulting in inflated confidence in one resolved tree or another and an increasing risk of false-positive inferences. Here we show that this is not the case; existing Bayesian methods do not lead to an inflation of statistical confidence, provided the evolutionary model is correct and uninformative priors are assumed. Posterior probabilities do not become increasingly unpredictable with increasing sequence length, and they exhibit conservative type I error rates, leading to a low rate of false-positive inferences. With infinite data, posterior probabilities give equal support for all resolved trees, and the rate of false inferences falls to zero. We conclude that there is no star tree paradox caused by not sampling unresolved trees.     

Mate choice for genetic quality when environments vary: suggestions for empirical progress

Mate choice for good-genes remains one of the most controversial evolutionary processes ever proposed. This is partly because strong directional choice should theoretically deplete the genetic variation that explains the evolution of this type of female mating preference (the so-called lek paradox). Moreover, good-genes benefits are generally assumed to be too small to outweigh opposing direct selection on females. Here, we review recent progress in the study of mate choice for genetic quality, focussing particularly on the potential for genotype by environment interactions (GEIs) to rescue additive genetic variation for quality, and thereby resolve the lek paradox. We raise five questions that we think will stimulate empirical progress in this field, and suggest directions for research in each area: (1) How is condition-dependence affected by environmental variation? (2) How important are GEIs for maintaining additive genetic variance in condition? (3) How much do GEIs reduce the signalling value of male condition? (4) How does GEI affect the multivariate version of the lek paradox? (5) Have mating biases for high-condition males evolved because of indirect benefits?     

The Role of Self-Sacrifice in Moral Dilemmas

Centuries’ worth of cultural stories suggest that self-sacrifice may be a cornerstone of our moral concepts, yet this notion is largely absent from recent theories in moral psychology. For instance, in the footbridge version of the well-known trolley car problem the only way to save five people from a runaway trolley is to push a single man on the tracks. It is explicitly specified that the bystander cannot sacrifice himself because his weight is insufficient to stop the trolley. But imagine if this were not the case. Would people rather sacrifice themselves than push another? In Study 1, we find that people approve of self-sacrifice more than directly harming another person to achieve the same outcome. In Studies 2 and 3, we demonstrate that the effect is not broadly about sensitivity to self-cost, instead there is something unique about sacrificing the self. Important theoretical implications about agent-relativity and the role of causality in moral judgments are discussed.     

Senescence as an adaptation to limit the spread of disease

Aging has the hallmarks of an evolved adaptation. It is controlled by genes that have been conserved over vast evolutionary distances, and most organisms are able to forestall aging in the most challenging of environments. But fundamental theoretical considerations imply that there can be no direct selection for aging. Senescence reduces individual fitness, and any group benefits are weak and widely dispersed over non-relatives. We offer a resolution to this paradox, suggesting a general mechanism by which senescence might have evolved as an adaptation. The proposed benefit is that senescence protects against infectious epidemics by controlling population density and increasing diversity of the host population. This mechanism is, in fact, already well-accepted in another context: it is the Red Queen Hypothesis for the evolution of sex. We illustrate the hypothesis using a spatially explicit agent-based model in which disease transmission is sensitive to population density as well as homogeneity. We find that individual senescence provides crucial population-level advantages, helping to control both these risk factors. Strong population-level advantages to individual senescence can overcome the within-population disadvantage of senescence. We conclude that frequent local extinctions provide a mechanism by which senescence may be selected as a population-level adaptation in its own right, without assuming pleiotropic benefits to the individual.     

The Biology of Invasions: The Genetic Adaptation Paradox

One of the most relevant topics in the biology of invasion concerns the genetic changes that occur subsequent to a species invasion, an issue of particular focus among conservation biologists. Colonizing a novel environment presents a genetic challenge to invading species because such species surely have not experienced the selective pressures presented by the environment. Here we ask, by what mechanisms and processes do alien species genetically naïve to their new environment, become successful invaders? We attempt to resolve this paradox by considering the interplay between an invader’s ability to modify its new environment, and genetic modifications imposed by the new environment. We postulate that epigenetic adaptations, and adaptive mutations are likely play a role in enhancing invasion success.     

Why are grape/fresh wine anthocyanins so simple and why is it that red wine color lasts so long?

Vitis vinifera red berries are characterized by anthocyanins whose chemical structures are among the simplest encountered in higher plants. On the contrary, many plants, including orchids, petunias, red cabbage, elderberries, potatoes for instance, have developed very complicated anthocyanins featuring side-chains at the available positions of the aglycone skeleton. Such pigments were shown to possess bio-physico-chemical properties not to be seen with the grape common anthocyanins. Among beverages (water, tea, beer, wine, coffee, juices, milk), red wine is the only one whose organoleptic properties improve with time and this is called ageing. The grape/fresh red wine pigments, after a few months, disappear from the wine giving birth to new pigments resulting from the wine spontaneous chemistry allowing it to remain red for many years. What are the wine pigments and why are they so stable is the purpose of this mini-review. The structural simplicity of grape anthocyanins and the long lasting colour of red wine is another French paradox; we call it French paradox II.     

Understanding height-structured competition in forests: is there an R* for light?

Tree species differ from one another in, and display trade-offs among, a wide range of attributes, including canopy and understorey growth and mortality rates, fecundity, height and crown allometry, and crown transmissivity. But how does this variation affect the outcome of interspecific competition and hence community structure? We derive criteria for the outcome of competition among tree species competing for light, given their allometric and life-history parameters. These criteria are defined in terms of a new simple whole life-cycle measure of performance, which provides a simple way to organize and understand the many ways in which species differ. The general case, in which all parameters can differ between species, can produce coexistence, founder control or competitive exclusion: thus, competition for light need not be hierarchical as implied by R^* theory. The special case in which species differ only in crown transmissivity produces neutral dynamics. The special case in which species differ in all parameters except crown transmissivity gives hierarchical competition, where the equivalent of R^* is Zˆ*, the height at which trees enter the canopy in an equilibrium monoculture.     

A note on a possible mathematical approach to the theory of individual freedom

As suggested in previous publications, freedom may be defined quantitatively as a restriction upon the choice of a number of activities. If the choice is determined by maximizing the satisfaction function, it is suggested that freedom may be defined in terms of the satisfaction function. If an individual is isolated and no physical restrictions limit his choice of activities, he is free to choose any activity in an amount which maximizes his satisfaction. This isolated state may be considered therefore as that of maximum freedom. If the individual interacts with another, he will choose different amounts of his object of satisfaction depending on whether he behaves egoistically or altruistically. But in either case the value chosen will not maximize his satisfaction function considered alone. A simple analytical expression is suggested as a measure of freedom in this case, and some problems which arise from this suggestion are mentioned.     

Toward a Resolution of the Paradox of Aggressive Displays: I. Optimal Deceit in the Communication of Fighting Ability

One inference from game theory models of animal conflict is that adversaries should not inform one another about concealed components of their fighting ability. This poses a paradox for the customary ethological account of aggressive displays in that it is usually assumed that the primary function of such behavior is to make such information available. To resolve the paradox, I propose that the information in aggressive displays may not be strictly truthful, but may instead represent “optimal deceit,” a balance between the advantages of deceit or bluffing and the disadvantages of selecting for skepticism in the receiver. Numerical simulation of this model was performed to examine the effects of differences in fighting ability and in the risk of injury in an escalated conflict. The model converged on an equilibrium level of deceit, even when the receiver was ignorant of the average level of deception being employed.     

Conflicting preferences within females: sexual selection versus species recognition

Preferences for mates within and between species are often harmonious, as traits that females prefer are usually more developed in conspecifics than heterospecifics. This need not be the case, however. When it is not, conflict between these arenas of mate choice can be resolved if females attend to different cues for each task. But this raises the potential for correlations among preferences to limit the opportunity for these two processes to operate independently. Here, we show that, within individual female pygmy swordtails (Xiphophorus pygmaeus), directional preferences for conspicuous ornamentation are inversely associated with discrimination against a sympatric heterospecific, Xiphophorus cortezi. Thus, mate choice among and within species need not be separate, independent processes; instead, they can be mechanistically intertwined. As a consequence, different arenas of mate choice can constrain one another, even when females assess multiple cues.     

The paradox of invasion in birds: competitive superiority or ecological opportunism?

Why can alien species succeed in environments to which they have had no opportunity to adapt and even become more abundant than many native species? Ecological theory suggests two main possible answers for this paradox: competitive superiority of exotic species over native species and opportunistic use of ecological opportunities derived from human activities. We tested these hypotheses in birds combining field observations and experiments along gradients of urbanization in New South Wales (Australia). Five exotic species attained densities in the study area comparable to those of the most abundant native species, and hence provided a case for the invasion paradox. The success of these alien birds was not primarily associated with a competitive superiority over native species: the most successful invaders were smaller and less aggressive than their main native competitors, and were generally excluded from artificially created food patches where competition was high. More importantly, exotic birds were primarily restricted to urban environments, where the diversity and abundance of native species were low. This finding agrees with previous studies and indicates that exotic and native species rarely interact in nature. Observations and experiments in the field revealed that the few native species that exploit the most urbanized environments tended to be opportunistic foragers, adaptations that should facilitate survival in places where disturbances by humans are frequent and natural vegetation has been replaced by man-made structures. Successful invaders also shared these features, suggesting that their success is not a paradox but can be explained by their capacity to exploit ecological opportunities that most native species rarely use.     

Testing evolutionary hypotheses for DNA barcoding failure in willows

The goal of DNA barcoding is to enable the rapid identification of taxa from short diagnostic DNA sequence profiles. But how feasible is this objective when many evolutionary processes, such as hybridization and selective sweeps, cause alleles to be shared among related taxa? In this issue of Molecular Ecology, Percy et al. (2014) test the full suite of seven candidate plant barcoding loci in a broad geographic sample of willow species. They show exceptional plastid haplotype sharing between species across continents, with most taxa not possessing a unique barcode sequence. Using population genetic and molecular dating analyses, they implicate hybridization and selective sweeps, but not incomplete lineage sorting, as the historical processes causing widespread haplotype sharing among willow taxa. This study represents an exceptional case of how poorly barcoding can perform, and highlights methodological issues using universal organellar regions for species identification.     

Take up the challenge! Opportunities for evolution research from resolving conflict in integrative taxonomy

What's in a species? The multiple connotations of the question tend to lack simple answers, and not surprisingly so. For example, speciation is a gradual process. Can we say when exactly a child has become an adult? We have precocious youngsters and late bloomers, and often, adults are in some ways childish. There are many triggers for and routes to adolescence. All this holds for speciation, and delimiting species can therefore be a tricky task. Recently, the field of integrative taxonomy has emerged—species delimitation based on multiple sources of evidence. Given that we expect species to exhibit peculiarities in at least one or a few aspects, might it be their alleles of a gene, their morphology, chemistry, behaviour, ecology, reproductive compatibility, or whatever, investigating not just one but several of these aspects makes it more likely that we capture such peculiarities. If the same pattern is found multiply, we talk about agreement among disciplines, and species delimitation is easy. But what if different disciplines tell different stories? Such disagreement makes species delimitation more difficult but is also an opportunity for evolutionary biology (Schlick‐Steiner et al. 2010). In this issue of Molecular Ecology, Andújar et al. (2014) present a comprehensive integrative‐taxonomic case study of Mesocarabus ground beetles including nomenclatural consequences. They resolve extensive disagreement among disciplines by invoking evolutionary explanations, and the process of conflict resolution thus advances knowledge on species boundaries and evolutionary processes simultaneously.     

Heterochromatin: a meiotic matchmaker?

During meiosis, the pairing of chromosomes is crucial for a successful partitioning of the genetic material and its transmission into the developing gamete. The classical view of meiotic pairing involves recombination between homologues as an integral part of the pairing mechanism. But, in cases where no recombination occurs, how do chromosome partners find one another? And how do they pair up and then segregate appropriately? Recently, a combination of molecular genetics and fluorescence in situ hybridization appears to have provided an answer to these questions by demonstrating a crucial role for heterochromatin in chromosome pairing.     

Love or fear: Can punishment promote cooperation?

Cooperation is a paradox: Why should one perform a costly behavior only to increase the fitness of another? Human societies, in which individuals cooperate with genetically unrelated individuals on a considerably larger scale than most mammals do, are especially puzzling in this regard. Recently, the threat of punishment has been given substantial attention as one of the mechanisms that could help sustain human cooperation in such situations. Nevertheless, using punishment to explain cooperation only leads to further questions: Why spend precious resources to penalize free‐riders, especially if others can avoid this investment and cheaters can punish you back? Here, it is argued that current evidence supports punishment as an efficient means for the maintenance of cooperation, and that the gravity of proposed limitations of punishment for maintaining cooperation may have been overestimated in previous studies due to the features of experimental design. Most notably, the importance of factors as characteristic of human societies as reputation and language has been greatly neglected. Ironically, it was largely the combination of the two that enabled humans to shape costly punishment into numerous low‐cost and less detrimental strategies that clearly can promote human cooperation.     

What is the true structure of liganded haemoglobin?

Does the crystal structure of a protein accurately represent its structure in solution? Or does the crystallization process perturb the structure significantly? Although aware of the problem, most crystallographers would argue that the highly solvated and weakly held lattice in protein crystals is, in general, unlikely to shift ordered parts of the molecule. In the case of conformationally flexible proteins, however, there is the possibility that one form might be favoured over another. Several lines of evidence suggest that this might be the case for the crystal structure of liganded Hb, although conflicting data exist.