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Biomedical subjects

Hanna Kokko

Publications and source records attributed to Hanna Kokko.

At least 19 recordsLinked to original sources
PubMed2026

The mechanistic and evolutionary diversity of programmed DNA elimination.

Beyond somatic mutations, the genetic makeup of an organism is often assumed to remain constant across all cells or nuclei within individuals. However, some organisms exhibit programmed DNA elimination (PDE), whereby specific cell lineages lose DNA segments or whole chromosomes. Building evidence indicates that PDE occurs in a wide range of eukaryotes and is linked to diverse cellular processes including gene silencing, germline differentiation, genome defence and sex determination. Here we compare PDE across broad phylogenetic groups, highlighting the mechanistic diversity, overlooked plasticity in genome integrity and the considerable gaps in our understanding of why PDE emerged recurrently and is maintained across the Tree of Life.

Journal Article↗
PubMed2006

Cheating can stabilize cooperation in mutualisms.

Mutualisms present a challenge for evolutionary theory. How is cooperation maintained in the face of selection for selfishness and cheating? Both theory and data suggest that partner choice, where one species preferentially directs aid to the more cooperative members of the other species, is central to cooperation in many mutualisms. However, the theory has only so far considered the evolutionary effects of partner choice on one of the species in a mutualism in isolation. Here, we investigate the co-evolution of cooperation and choice in a choosy host and its symbiont. Our model reveals that even though choice and cooperation may be initially selected, it will often be unstable. This is because choice reduces variation in the symbiont and, therefore, tends to remove the selective incentive for its own maintenance (a scenario paralleled in the lek paradox in female choice and policing in within-species cooperation). However, we also show that when variability is reintroduced into symbionts each generation, in the form of less cooperative individuals, choice is maintained. This suggests that the presence of cheaters and cheater species in many mutualisms is central to the maintenance of partner choice and, paradoxically, cooperation itself.

Animals↗
PubMed2006

Mate choice evolution, dominance effects, and the maintenance of genetic variation.

Female mate choice influences the maintenance of genetic variation by altering the mating success of males with different genotypes. The evolution of preferences themselves, on the other hand, depends on genetic variation present in the population. Few models have tracked this feedback between a choice gene and its effects on genetic variation, in particular when genes that determine offspring viability and attractiveness have dominance effects. Here we build a population genetic model that allows comparing the evolution of various choice rules in a single framework. We first consider preferences for good genes and show that focused preferences for homozygotes evolve more easily than broad preferences, which allow heterozygous males high mating success too. This occurs despite better maintenance of genetic diversity in the latter scenario, and we discuss why empirical findings of superior mating success of heterozygous males consequently do not immediately lead to a better understanding of the lek paradox. Our results thus suggest that the mechanisms that help maintain genetic diversity also have a flipside of making female choice an inaccurate means of producing the desired kind of offspring. We then consider preferences for heterozygosity per se, and show that these evolve only under very special conditions. Choice for compatible genotypes can evolve but its selective advantage diminishes quickly due to frequency-dependent selection. Finally, we show that our model reproduces earlier results on selfing, when the female choice strategy produces assortative mating. Overall, our model indicates that various forms of heterozygote-favouring (or variable) female choice pose a problem for the theory of sexual ornamentation based on indirect benefits, rather than a solution.

Animals↗
PubMed2006

From individual dispersal to species ranges: perspectives for a changing world.

Dispersal is often risky to the individual, yet the long-term survival of populations depends on having a sufficient number of individuals that move, find each other, and locate suitable breeding habitats. This tension has consequences that rarely meet our conservation or management goals. This is particularly true in changing environments, which makes the study of dispersal urgently topical in a world plagued with habitat loss, climate change, and species introductions. Despite the difficulty of tracking mobile individuals over potentially vast ranges, recent research has revealed a multitude of ways in which dispersal evolution can either constrain, or accelerate, species' responses to environmental changes.

Adaptation, Biological↗
PubMed2006

From hawks and doves to self-consistent games of territorial behavior.

Explaining the "prior-residence effect" (automatic owner status of individuals who arrived first in an area) was one of the very first applications of game theory in animal behavior. These models, however, predict paradoxical solutions where intruders always win, with no satisfactory explanation for the absence of such cases in nature. We propose a solution based on new developments in evolutionary game theory. A self-consistent model with feedbacks between individual behavior and population dynamics produces qualitatively different frequency-dependent selection on intruders (floaters) than on territory owners. Starting with an ancestral population with no respect for ownership, the most likely evolutionary end point is complete or partial respect. Conventional rules of conflict resolution thus can rely on "uncorrelated asymmetries" without differences in resource-holding power or territory value, although they will be strengthened by such differences. We also review the empirical literature on animal contests, testing whether asymmetries in resource-holding power are required to explain the observations. Despite much empirical effort, results remain inconclusive, because experiments are often unable to distinguish between the motivation of individuals to fight and the behavioral outcome of a contest. To help arrive at conclusive answers, we suggest a standardized empirical approach to quantify prior-residence effects.

Animals↗
PubMed2006

Sex, death and tragedy.

The population consequences of sexual conflict are relatively unexplored. In a recent paper, Le Galliard et al. now show that males of the common lizard Lacerta vivipara cause such damage to females that male-biased populations decrease in size, posing a real risk to the persistence of local lizard populations. Their study reveals surprising parallels between sexual conflict and the tragedy of the commons, where selfish competition over females destroys the very resource (i.e. the females) over which the males are fighting.

Aggression↗
PubMed2006

Lonely hearts or sex in the city? Density-dependent effects in mating systems.

Two very basic ideas in sexual selection are heavily influenced by numbers of potential mates: the evolution of anisogamy, leading to sex role differentiation, and the frequency dependence of reproductive success that tends to equalize primary sex ratios. However, being explicit about the numbers of potential mates is not typical to most evolutionary theory of sexual selection. Here, we argue that this may prevent us from finding the appropriate ecological equilibria that determine the evolutionary endpoints of selection. We review both theoretical and empirical advances on how population density may influence aspects of mating systems such as intrasexual competition, female choice or resistance, and parental care. Density can have strong effects on selective pressures, whether or not there is phenotypic plasticity in individual strategies with respect to density. Mating skew may either increase or decrease with density, which may be aided or counteracted by changes in female behaviour. Switchpoints between alternative mating strategies can be density dependent, and mate encounter rates may influence mate choice (including mutual mate choice), multiple mating, female resistance to male mating attempts, mate searching, mate guarding, parental care, and the probability of divorce. Considering density-dependent selection may be essential for understanding how populations can persist at all despite sexual conflict, but simple models seem to fail to predict the diversity of observed responses in nature. This highlights the importance of considering the interaction between mating systems and population dynamics, and we strongly encourage further work in this area.

Animals↗
PubMed2006

Why do female migratory birds arrive later than males?

1. In migratory birds males tend to arrive first on breeding grounds, except in sex-role reversed species. The two most common explanations are the rank advantage hypothesis, in which male-male competition for breeding sites drives stronger selection for early arrival in males than females, and the mate opportunity hypothesis, which relies on sexual selection, as early arrival improves prospects of mate acquisition more for males than for females. 2. To date, theoretical work has focused on selection for early arrival within a single sex, usually male. However, if fitness depends on territory quality, selection for early arrival should operate on both sexes. Here we use two independent modelling approaches to explore the evolution of protandry (male-first arrival) and protogyny (female-first arrival) under the rank advantage and mate opportunity hypotheses. 3. The rank advantage hypothesis, when operating alone, fails to produce consistent patterns of protandry, despite our assumption that males must occupy territories before females. This is because an individual of either sex benefits if it out-competes same-sex competitors. Rather than promoting protandry, the rank advantage mechanism can sometimes result in protogyny. Female-female competition is stronger than male-male competition early in the season, if females compete for a resource (territories occupied by males) that is initially less common than the resource of interest to males (unoccupied territories). 4. Our results support the mate opportunity hypothesis as an explanation of why protandry is the norm in migratory systems. Male-biased adult sex ratios and high levels of sperm competition (modelled as extra-pair young: EPY) both produce protandry as a result of sexual selection. Protogyny is only observed in our models with female-biased sex ratios and low EPY production. 5. We also show that the effects of sex ratio biases are much stronger than those of EPY production, explore the evidence for sex ratio biases and extra-pair paternity in migratory species and suggest future research directions.

Animal Migration↗
PubMed2006

Can non-directional male mating preferences facilitate honest female ornamentation?

Recent studies have demonstrated male mate choice for female ornaments in species without sex-role reversal. Despite these empirical findings, little is known about the adaptive dynamics of female signalling, in particular the evolution of male mating preferences. The evolution of traits that signal mate quality is more complex in females than in males because females usually provide the bulk of resources for the developing offspring. Here, we investigate the evolution of male mating preferences using a mathematical model which: (i) specifically accounts for the fact that females must trade-off resources invested in ornaments with reproduction; and (ii) allows male mating preferences to evolve a non-directional shape. The optimal adaptive strategy for males is to develop stabilizing mating preferences for female display traits to avoid females that either invests too many or too few resources in ornamentation. However, the evolutionary stability of this prediction is dependent upon the level of error made by females when allocating resources to either signal or fecundity.

Animals↗
PubMed2006

When not to avoid inbreeding.

Avoidance of incestuous matings is widely reported across many animal taxa, and the adaptive value of such behavior is explained through inbreeding depression. However, an old and somewhat neglected theoretical result predicts that inbred matings offer another, positive effect on the inclusive fitness of parents: an individual who mates with a relative will help that relative to spread genes identical by descent. This benefit can be substantial, if the additional mating achieved by the relative does not harm his mating success otherwise, and in the context of selfing in plants the phenomenon is well known. Here, we develop a model that derives expected values of inbreeding tolerance, that is, the magnitude of inbreeding depression that is required to make individuals avoid inbreeding, for different animal life histories and parental investment patterns. We also distinguish between simultaneous and sequential mate choice, and show that inbreeding tolerance should often be remarkably high in the latter scenario in particular, although egalitarian parental care will lead to lower tolerance. There is a mismatch between theory and data: the almost complete lack of cases where individuals prefer to mate incestuously is at odds with a large overlap between the predicted range of inbreeding tolerance and estimates of inbreeding depression found in nature. We discuss four different solutions to this enigma, and suggest that inbreeding tolerance, where it is found, should not always be attributed to a simple constraint that has prevented finding any other mate.

Animals↗
PubMed2005

Territorial defense, territory size, and population regulation.

The carrying capacity of an environment is determined partly by how individuals compete over the available resources. To territorial animals, space is an important resource, leading to conflict over its use. We build a model where the carrying capacity for an organism in a given environment results from the evolution of territorial defense effort and the consequent space use. The same evolutionary process can yield two completely different modes of population regulation. Density dependence arises through expanding and shrinking territories if fecundity is low, breeding success increases gradually with territory size, and/or defense is cheap. By contrast, when fecundity is high, breeding success sharply saturates with territory size, and/or defense is costly, we predict fixed territory sizes and regulation by floaters. These "surplus" individuals form a buffer against population fluctuations. Yet floaters can also harm breeder performance, and by comparing population growth of a territorial population to a nonterritorial (and individually suboptimal) alternative, we can quantify the harmful effect of evolutionary conflict on population performance. Territoriality has often been found to increase population stability, but this may come at a cost of reduced equilibrium densities.

Animals↗
PubMed2005

Sexual selection when fertilization is not guaranteed.

Much of the theory of sexual selection assumes that females do not generally experience difficulties getting their eggs fertilized, yet sperm limitation is occasionally documented. How often does male limitation form a selection for female traits that improve their mating rate? The question is difficult to test, because if such traits evolve to be efficient, sperm limitation will no longer appear to be a problem to females. Here, we suggest that changes in choosiness between populations, and in particular between virgin and mated females, offer an efficient way to test this hypothesis. We model the "wallflower effect," that is, changes in female preferences due to time and mortality costs of remaining unmated (for at least some time). We show that these costs cause adaptive reductions of female choice, even if mate encounter rates appear high and females only rarely end their lives unfertilized. We also consider the population consequences of plastic or fixed mate preferences at different mate encounter rates. If mate choice is plastic, we confirm earlier verbal models that virgins should mate relatively indiscriminately, but plastic increase of choosiness in later matings can compensate and intensify sexual selection on the male trait, particularly if there is last male sperm precedence. Plastic populations will cope well with unusual conditions: eagerness of virgins leads to high reproductive output and a relaxation of sexual selection at low population densities. If females lack such plasticity, however, population-wide reproductive output may be severely reduced, whereas sexual selection on male traits remains strong.

Animals↗
PubMed2004

Can too strong female choice deteriorate male ornamentation?

Competition for limited resources can have fundamental implications for population dynamics. However, the effects of resource depletion have rarely been discussed in the context of sexual selection, even though mate choice typically favours males who outperform others in securing access to some limited resource. Here, we develop a model to investigate the question of resource competition as a form of male-male competition in the context of male sexual displays. We phrase our model in terms of male bowerbirds either searching for or stealing resources (ornamental objects) valued by females, and compare the model findings with published studies of time allocation to various activities in different species of bowerbirds. The basic idea of the model, however, extends to cases where the resource is used less directly for the development of sexual ornamentation, such as males excluding others' access to food. We show that if males compete for resources used in sexual displays, intense female preference for high-quality displays can lead to poorer prospects for efficient choice by females. This is because males benefit from excluding others' access to resources used in displays, damaging the overall efficiency of resource use in the population, and the accuracy with which females can judge male ability to gain such resources. The evolution of female choice may therefore have a self-limiting nature when it poses a selection pressure on male resource acquisition.

Animals↗
PubMed2004

Priority versus brute force: when should males begin guarding resources?

When should males begin guarding a resource when both resources and guarders vary in quality? This general problem applies, for example, to migrant birds occupying territories in the spring and to precopula in crustaceans where males grab females before they molt and become receptive. Previous work has produced conflicting predictions. Theory on migrant birds predicts that the strongest competitors should often arrive first, whereas some models of mate guarding have predicted that the strongest competitors wait and then simply usurp a female from a weaker competitor. We build a general model of resource guarding that allows varying the ease with which takeovers occur. The model is phrased in terms of mate-guarding crustaceans, but the same logic can be applied to other forms of resource acquisition where priority plays a role but takeovers might be possible too. The race to secure breeding positions can lead to strong competitors (large males) taking females earliest, even though this means accepting a lower-quality female. Paradoxically, this means that small males, which have fewer breeding opportunities, are more choosy than larger ones. Such solutions are found when takeovers are impossible. The easier the takeovers and the higher the rate of finding guarded resources, the more likely are solutions where guarding durations are short, where strong competitors begin guarding only just before breeding, and where they do this by usurping the resource. The relationship between an individual's competitive ability and its timing of resource acquisition can also be nonlinear if takeovers are moderately common; if this is the case, then males of intermediate size guard the longest.

Animals↗
PubMed2003

Quantifying male attractiveness.

Genetic models of sexual selection are concerned with a dynamic process in which female preference and male trait values coevolve. We present a rigorous method for characterizing evolutionary endpoints of this process in phenotypic terms. In our phenotypic characterization the mate-choice strategy of female population members determines how attractive females should find each male, and a population is evolutionarily stable if population members are actually behaving in this way. This provides a justification of phenotypic explanations of sexual selection and the insights into sexual selection that they provide. Furthermore, the phenotypic approach also has enormous advantages over a genetic approach when computing evolutionarily stable mate-choice strategies, especially when strategies are allowed to be complex time-dependent preference rules. For simplicity and clarity our analysis deals with haploid mate-choice genetics and a male trait that is inherited phenotypically, for example by vertical cultural transmission. The method is, however, easily extendible to other cases. An example illustrates that the sexy son phenomenon can occur when there is phenotypic inheritance of the male trait.

Animals↗
PubMed2003

The evolution of mate choice and mating biases.

We review the current status of three well-established models (direct benefits, indirect benefits and sensory drive) and one newcomer (antagonistic chase-away) of the evolution of mate choice and the biases that are expressed during choice. We highlight the differences and commonalities in the underlying genetics and evolutionary dynamics of these models. We then argue that progress in understanding the evolution of mate choice is currently hampered by spurious distinctions among models and a misguided tendency to test the processes underlying each model as mutually exclusive alternatives. Finally, we suggest potentially fruitful directions for future theoretical and empirical research.

Animals↗