Filial cannibalism of Nabis pseudoferus is not evolutionarily optimal foraging strategy.

Using a recursion model with real parameters of Nabis pseudoferus, we show that its filial cannibalism is an optimal foraging strategy for life reproductive success, but it is not an evolutionarily optimal foraging strategy, since it cannot maximize the descendant's number at the end of the reproductive season. Cannibalism is evolutionarily rational, when the number of newborn offspring produced from the cannibalized offspring can compensate the following two effects: (a) The cannibalistic lineage wastes time, since the individuals hatched from eggs produced by cannibalism start to reproduce later. (b) Cannibalism eliminates not only one offspring, but also all potential descendants from the cannibalized offspring during the rest of reproductive season. In our laboratory trials, from conspecific prey Nabis pseudoferus did not produce newborn nymphs enough to compensate the above two effects.

Genesis of ectosymbiotic features based on commensalistic syntrophy.

The symbiogenetic origin of eukaryotes with mitochondria is considered a major evolutionary transition. The initial interactions and conditions of symbiosis, along with the phylogenetic affinity of the host, are widely debated. Here, we focus on a possible evolutionary path toward an association of individuals of two species based on unidirectional syntrophy. With the backing of a theoretical model, we hypothesize that the first step in the evolution of such symbiosis could be the appearance of a linking structure on the symbiont's membrane, using which it forms an ectocommensalism with its host. We consider a commensalistic model based on the syntrophy hypothesis in the framework of coevolutionary dynamics and mutant invasion into a monomorphic resident system (evolutionary substitution). We investigate the ecological and evolutionary stability of the consortium (or symbiotic merger), with vertical transmissions playing a crucial role. The impact of the 'effectiveness of vertical transmission' on the dynamics is also analyzed. We find that the transmission of symbionts and the additional costs incurred by the mutant determine the conditions of fixation of the consortia. Additionally, we observe that small and highly metabolically active symbionts are likely to form the consortia.

Coevolutionary stability of host-symbiont systems with mixed-mode transmission.

The coevolution of hosts and symbionts based on virulence and mode of transmission is a complex and diverse biological phenomenon. We introduced a conceptual model to study the stable coexistence and coevolution of an obligate symbiont (mutualist or parasite) with mixed-mode transmission and its host. Using an age-structured Leslie model for the host, we demonstrated how the obligate symbiont could modify the host's life history traits (survival and fecundity) and the long-term growth rate of the infected lineage. When the symbiont is vertically transmitted, we found that the host and its symbiont could maximize the infected lineage's evolutionary success (multi-level selection). Our model showed that symbionts' effect on host longevity and reproduction might differ, even be opposing, and their net effect might often be counterintuitive. The evolutionary stability of the ecologically stable coexistence was analyzed in the framework of coevolutionary dynamics. Moreover, we found conditions for the ecological and evolutionary stability of the resident host-symbiont pair, which does not allow invasion by rare mutants (each mutant dies out by ecological selection). We concluded that, within the context of our simplified model conditions, a host-symbiont system with mixed-mode transmission is evolutionarily stable unconditionally only if the host can maximize the Malthusian parameters of the infected and non-infected lineages using the same strategy. Finally, we performed a game-theoretical analysis of our selection situation and compared two stability definitions.

Definitions of parasitism, considering its potentially opposing effects at different levels of hierarchical organization.

An annotated synthesis of textbook definitions of parasitism is presented. Most definitions declare parasitism is a long-lasting relationship between individuals of different species harming the hosts. The infection-induced costs are interpreted as diseases in the medical-veterinary literature. Alternatively, evolutionary ecologists interpret it as a reduction of host's fitness (longevity, fertility or both). Authors often assume that such effects decrease host population growth and select for antiparasitic defences, which is not necessarily true because infections may simultaneously express opposite effects at different levels of biological organization. (i) At the cellular level, infection-induced cell growth, longevity and multiplication may yield tumours maladaptive at higher levels. (ii) At the individual level, reduced host longevity, fertility or both are interpreted as disease symptoms or reduced fitness. (iii) Contrary to common sense, the growth rate of infected host lineages may increase in parallel with the individuals' reduced survival and fertility. This is because selection favours not only the production of more offspring but also their faster production. (iv) Finally, infections that reduce host individuals' or lineages' fitness may still increase infected host populations' growth rate in the context of ecological competition. Therefore, differences between parasitism and mutualism may depend on which level of organization one focuses on.

Subsistence of sib altruism in different mating systems and Haldane's arithmetic.

The moral rule "Risk your life to save your family members" is, at the same time, a biological phenomenon. The prominent population geneticist, J.B.S. Haldane told his friends that he would risk his life to save two drowning brothers, but not one - so the story goes. In biological terms, Haldane's arithmetic claims that sib altruism is evolutionarily rational, whenever by "self-sacrifice" an altruistic gene "rescues", on average, more than one copy of itself in its lineage. Here, we derive conditions for evolutionary stability of sib altruism, using population genetic models for three mating systems (monogamy, promiscuity and polygyny) with linear and non-linear group effect on the siblings' survival rate. We show that for all considered selection situations, the condition of evolutionary stability is equivalent to Haldane's arithmetic. The condition for evolutionary stability is formulated in terms of genetic relatedness and the group effect on the survival probability, similarly to the classical Hamilton's rule. We can set up a "scale of mating systems", since in pairwise interactions the chance of evolutionary stability of sib altruism decreases in this order: monogamy, polygyny and promiscuity. Practice of marrying and siblings' solidarity are moral rules in a secular world and in various religious traditions. These moral rules are not evolutionarily independent, in the sense that the subsistence of sib altruism is more likely in a monogamous population.

State-controlled epidemic in a game against a novel pathogen.

The pandemic reminded us that the pathogen evolution still has a serious effect on human societies. States, however, can prepare themselves for the emergence of a novel pathogen with unknown characteristics by analysing potential scenarios. Game theory offers such an appropriate tool. In our game-theoretical framework, the state is playing against a pathogen by introducing non-pharmaceutical interventions to fulfil its socio-political goals, such as guaranteeing hospital care to all needed patients, keeping the country functioning, while the applied social restrictions should be as soft as possible. With the inclusion of activity and economic sector dependent transmission rate, optimal control of lockdowns and health care capacity management is calculated. We identify the presence and length of a pre-symptomatic infectious stage of the disease to have the greatest effect on the probability to cause a pandemic. Here we show that contrary to intuition, the state should not strive for the great expansion of its health care capacities even if its goal is to provide care for all requiring it and minimize the cost of lockdowns.

Best Reply Player Against Mixed Evolutionarily Stable Strategy User.

We consider matrix games with two phenotypes (players): one following a mixed evolutionarily stable strategy and another one that always plays a best reply against the action played by its opponent in the previous round (best reply player, BR). We focus on iterated games and well-mixed games with repetition (that is, the mean number of repetitions is positive, but not infinite). In both interaction schemes, there are conditions on the payoff matrix guaranteeing that the best reply player can replace the mixed ESS player. This is possible because best reply players in pairs, individually following their own selfish strategies, develop cycles where the bigger payoff can compensate their disadvantage compared with the ESS players. Well-mixed interaction is one of the basic assumptions of classical evolutionary matrix game theory. However, if the players repeat the game with certain probability, then they can react to their opponents' behavior. Our main result is that the classical mixed ESS loses its general stability in the well-mixed population games with repetition in the sense that it can happen to be overrun by the BR player.

When optimal foragers meet in a game theoretical conflict: A model of kleptoparasitism.

Kleptoparasitism can be considered as a game theoretical problem and a foraging tactic at the same time, so the aim of this paper is to combine the basic ideas of two research lines: evolutionary game theory and optimal foraging theory. To unify these theories, firstly, we take into account the fact that kleptoparasitism between foragers has two consequences: the interaction takes time and affects the net energy intake of both contestants. This phenomenon is modeled by a matrix game under time constraints. Secondly, we also give freedom to each forager to avoid interactions, since in optimal foraging theory foragers can ignore each food type (we have two prey types: either a prey item in possession of another predator or a free prey individual is discovered). The main question of the present paper is whether the zero-one rule of optimal foraging theory (always or never select a prey type) is valid or not, in the case where foragers interact with each other? In our foraging game we consider predators who engage in contests (contestants) and those who never do (avoiders), and in general those who play a mixture of the two strategies. Here the classical zero-one rule does not hold. Firstly, the pure avoider phenotype is never an ESS. Secondly, the pure contestant can be a strict ESS, but we show this is not necessarily so. Thirdly, we give an example when there is mixed ESS.

Do Development and Diet Determine the Degree of Cannibalism in Insects? To Eat or Not to Eat Conspecifics.

Cannibalism in insects plays an important role in ecological relationships. Nonetheless, it has not been studied as extensively as in other arthropods groups (e.g., Arachnida). From a theoretical point of view, cannibalism has an impact on the development of more realistic stage-structure mathematical models. Additionally, it has a practical application for biological pest control, both in mass-rearing and out in the field through inoculative releases. In this paper, the cannibalistic behavior of two species of predatory bugs was studied under laboratory conditions-one of them a generalist predator (strictly carnivorous), Nabis pseudoferus, and the other a true omnivore (zoophytophagous), Nesidiocoris tenuis-and compared with the intraguild predation (IGP) behavior. The results showed that cannibalism in N. pseudoferus was prevalent in all the developmental stages studied, whereas in N. tenuis, cannibalism was rarely observed, and it was restricted mainly to the first three nymphal stages. Cannibalism and intraguild predation had no linear relationship with the different cannibal-prey size ratios, as evaluated by the mortality rates and survival times, although there were variations in cannibalism between stages, especially for N. pseudoferus. The mathematical model's implications are presented and discussed.

A temporal model of territorial defence with antagonistic interactions.

Territorial behaviour is an important part of the lives of many animals. Once a territory has been acquired, an animal may spend its entire life on it, and may have to repeatedly defend it from conspecifics. Some species make great investments in the defence of a territory, and this defence can be costly, in terms of time, energy and risk of injury. Time costs in particular have rarely been explicitly factored into such models. In this paper we consider a model of territorial defence which includes both population dynamic and time delay elements, building upon recent advances in time constraint models. Populations may divide into two distinct types, where one type makes no effort to control territories. We shall call this type nomads, and the other type territorials. Here the territory owners must divide their time between patrolling and foraging, and this balance is their only strategic decision. We show how to find the evolutionarily stable patrolling strategy and the population composition of territorials and nomads, and consider some examples demonstrating key situations. We see that both time constraints and population density pressure are crucial to influencing behaviour. In particular we find cases with both territorial individuals and nomads where a mixed, either pure or both pure patrolling strategies are evolutionarily stable. In different conditions either nomads or territorials can be absent, and indeed for a significant range of parameter combinations the population can exhibit tristability, with three distinct ecologically stable population compositions: with both nomads and territorials, only nomads or only territorials.

The ESS for evolutionary matrix games under time constraints and its relationship with the asymptotically stable rest point of the replicator dynamics.

Recently we interpreted the notion of ESS for matrix games under time constraints and investigated the corresponding state in the polymorphic situation. Now we give two further static (monomorphic) characterizations which are the appropriate analogues of those known for classical evolutionary matrix games. Namely, it is verified that an ESS can be described as a neighbourhood invader strategy independently of the dimension of the strategy space in our non-linear situation too, that is, a strategy is an ESS if and only if it is able to invade and completely replace any monomorphic population which totally consists of individuals following a strategy close to the ESS. With the neighbourhood invader property at hand, we establish a dynamic characterization under the replicator dynamics in two dimensions which corresponds to the strong stability concept for classical evolutionary matrix games. Besides, in some special cases, we also prove the stability of the corresponding rest point in higher dimensions.

To save or not to save your family member's life? Evolutionary stability of self-sacrificing life history strategy in monogamous sexual populations.

BACKGROUND: For the understanding of human nature, the evolutionary roots of human moral behaviour are a key precondition. Our question is as follows: Can the altruistic moral rule "Risk your life to save your family members, if you want them to save your life" be evolutionary stable? There are three research approaches to investigate this problem: kin selection, group selection and population genetics modelling. The present study is strictly based on the last approach. RESULTS: We consider monogamous and exogamous families, where at an autosomal locus, dominant-recessive alleles determine the phenotypes in a sexual population. Since all individuals' survival rate is determined by their altruistic family members, we introduce a new population genetics model based on the mating table approach and adapt the verbal definition of evolutionary stability to genotypes. In general, when the resident is recessive, a homozygote is an evolutionarily stable genotype (ESG), if the number of survivors of the resident genotype of the resident homozygote family is greater than that of non-resident heterozygote survivors of the family of the resident homozygote and mutant heterozygote genotypes. Using the introduced genotype dynamics we proved that in the recessive case ESG implies local stability of the altruistic genotype. We apply our general ESG conditions for self-sacrificing life history strategy when the number of new-born offspring does not depend on interactions within the family and the interactions are additive. We find that in this case our ESG conditions give back Hamilton's rule for evolutionary stability of the self-sacrificing life history strategy. CONCLUSIONS: In spite of the fact that the kidney transplantations was not a selection factor during the earlier human evolution, nowadays "self-sacrificing" can be observed in the live donor kidney transplantations, when the donor is one of the family members. It seems that selection for self-sacrificing in family produced an innate moral tendency in modulating social cognition in human brain.

Juvenile honest food solicitation and parental investment as a life history strategy: A kin demographic selection model.

Parent-offspring communication remains an unresolved challenge for biologist. The difficulty of the challenge comes from the fact that it is a multifaceted problem with connections to life-history evolution, parent-offspring conflict, kin selection and signalling. Previous efforts mainly focused on modelling resource allocation at the expense of the dynamic interaction during a reproductive season. Here we present a two-stage model of begging where the first stage models the interaction between nestlings and parents within a nest and the second stage models the life-history trade-offs. We show in an asexual population that honest begging results in decreased variance of collected food between siblings, which leads to mean number of surviving offspring. Thus, honest begging can be seen as a special bet-hedging against informational uncertainty, which not just decreases the variance of fitness but also increases the arithmetic mean.

Opportunistic random searcher versus intentional search image user.

We consider two types of optimal foragers: a random searcher and a search image user. A search image user can find its desired prey with higher and undesired prey with lower probability than a random searcher. Our model considers the density-dependent travelling time and the time duration of reproduction (oviposition). In the framework of optimal foraging theory for one predator-two prey systems, we find that there are ranges of prey densities in which the search image user has a higher net energy intake, and there are other ranges of prey densities in which the random searcher has higher net energy intake. The damsel bug Nabis pseudoferus Remane (Hemiptera: Nabidae) is a generalist predator rather than an omnivore. This species has a wide range of arthropod prey (predominantly insects and mites). Several aspects of the biology of this species have been studied, especially its cannibalistic behaviour, which is a quite important feature because N. pseudoferus is often used as a biological control agent against lepidopteran pests in greenhouse crops. Experimentally, we found that Nabis is a search image user in the above sense.

The ESS and replicator equation in matrix games under time constraints.

Recently, we introduced the class of matrix games under time constraints and characterized the concept of (monomorphic) evolutionarily stable strategy (ESS) in them. We are now interested in how the ESS is related to the existence and stability of equilibria for polymorphic populations. We point out that, although the ESS may no longer be a polymorphic equilibrium, there is a connection between them. Specifically, the polymorphic state at which the average strategy of the active individuals in the population is equal to the ESS is an equilibrium of the polymorphic model. Moreover, in the case when there are only two pure strategies, a polymorphic equilibrium is locally asymptotically stable under the replicator equation for the pure-strategy polymorphic model if and only if it corresponds to an ESS. Finally, we prove that a strict Nash equilibrium is a pure-strategy ESS that is a locally asymptotically stable equilibrium of the replicator equation in n-strategy time-constrained matrix games.

Survival phenotype, selfish individual versus Darwinian phenotype.

Consider and infinitely large asexual population without mutations and direct interactions. The activities of an individual determine the fecundity and the survival probability of individuals, moreover each activity takes time. We view this population model as a simple combination of life history and optimal foraging models. The phenotypes are given by probability distributions on these activities. We concentrate on the following phenotypes defined by optimization of different objective functions: selfish individual (maximizes the average offspring number during life span), survival phenotype (maximizes the probability of non-extinction of descendants) and Darwinian phenotype (maximizes the phenotypic growth rate). We find that the objective functions above can achieve their maximum at different activity distributions, in general. We find that the objective functions above can achieve their maximum at different activity distributions, in general. The novelty of our work is that we let natural selection act on the different objective functions. Using the classical Darwinian reasoning, we show that in our selection model the Darwinian phenotype outperforms all other phenotypes.

Evolutionary stability for matrix games under time constraints.

Game theory focuses on payoffs and typically ignores time constraints that play an important role in evolutionary processes where the repetition of games can depend on the strategies, too. We introduce a matrix game under time constraints, where each pairwise interaction has two consequences: both players receive a payoff and they cannot play the next game for a specified time duration. Thus our model is defined by two matrices: a payoff matrix and an average time duration matrix. Maynard Smith's concept of evolutionary stability is extended to this class of games. We illustrate the effect of time constraints by the well-known prisoner's dilemma game, where additional time constraints can ensure the existence of unique evolutionary stable strategies (ESS), both pure and mixed, or the coexistence of two pure ESS. Our general results may be useful in several fields of biology where evolutionary game theory is applied, principally in ecological games, where time constraints play an inevitable role.

A two-agent model applied to the biological control of the sugarcane borer (Diatraea saccharalis) by the egg parasitoid Trichogramma galloi and the larvae parasitoid Cotesia flavipes.

The paper is aimed at a methodological development in biological pest control. The considered one pest two-agent system is modelled as a verticum-type system. Originally, linear verticum-type systems were introduced by one of the authors for modelling certain industrial systems. These systems are hierarchically composed of linear subsystems such that a part of the state variables of each subsystem affect the dynamics of the next subsystem. Recently, verticum-type system models have been applied to population ecology as well, which required the extension of the concept a verticum-type system to the nonlinear case. In the present paper the general concepts and technics of nonlinear verticum-type control systems are used to obtain biological control strategies in a two-agent system. For the illustration of this verticum-type control, these tools of mathematical systems theory are applied to a dynamic model of interactions between the egg and larvae populations of the sugarcane borer (Diatraea saccharalis) and its parasitoids: the egg parasitoid Trichogramma galloi and the larvae parasitoid Cotesia flavipes. In this application a key role is played by the concept of controllability, which means that it is possible to steer the system to an equilibrium in given time. In addition to a usual linearization, the basic idea is a decomposition of the control of the whole system into the control of the subsystems, making use of the verticum structure of the population system. The main aim of this study is to show several advantages of the verticum (or decomposition) approach over the classical control theoretical model (without decomposition). For example, in the case of verticum control the pest larval density decreases below the critical threshold value much quicker than without decomposition. Furthermore, it is also shown that the verticum approach may be better even in terms of cost effectiveness. The presented optimal control methodology also turned out to be an efficient tool for the "in silico" analysis of the cost-effectiveness of different biocontrol strategies, e.g. by answering the question how far it is cost-effective to speed up the reduction of the pest larvae density, or along which trajectory this reduction should be carried out.

Can Interactions Between an Omnivorous Hemipteran and an Egg Parasitoid Limit the Level of Biological Control for the Tomato Pinworm?

Relationships between the omnivorous predator Nesidiocoris tenuis (Reuter) and the egg parasitoid Trichogramma achaeae Nagaraja and Nagarkatti were studied in the laboratory (no-choice and choice assays, and functional responses) and in a greenhouse experiment. Both natural enemies are utilized in the biological control of tomato pinworm on greenhouse-grown tomato crops. Three different food items were offered to the predator: nonparasitized prey, prey parasitized for less than 4 d by T. achaeae, and prey parasitized for more than 4 d by the parasitoid. There were significant differences in consumption of food types, with highest consumption for nonparasitized prey, followed by parasitized (<4 d) and then parasitized (>4 d), both in no-choice and choice trials. At the same time, the predator causes a significant mortality in the prey (over 80%) regardless of previous parasitism, resulting in a very coincidental intraguild predation detrimental to the parasitoid. It has also been observed that there was a change in the functional response by the predator from Type II in presence of nonparasitized prey to Type I when there was a combination of parasitized and nonparasitized prey. This represents an increase of instantaneous search rate (a') and a decrease of handling time (Th), which indicates a change in feeding behavior on the two prey types. Under greenhouse conditions, the intraguild predation reduced the percentage of parasitism by T. achaeae in just over 20%. However, when both natural enemies were present, a better control of pest Tuta absoluta (Meyrick) was achieved than in the case of application of any of them alone.

Optimal forager against ideal free distributed prey.

The introduced dispersal-foraging game is a combination of prey habitat selection between two patch types and optimal-foraging approaches. Prey's patch preference and forager behavior determine the prey's survival rate. The forager's energy gain depends on local prey density in both types of exhaustible patches and on leaving time. We introduce two game-solution concepts. The static solution combines the ideal free distribution of the prey with optimal-foraging theory. The dynamical solution is given by a game dynamics describing the behavioral changes of prey and forager. We show (1) that each stable equilibrium dynamical solution is always a static solution, but not conversely; (2) that at an equilibrium dynamical solution, the forager can stabilize prey mixed patch use strategy in cases where ideal free distribution theory predicts that prey will use only one patch type; and (3) that when the equilibrium dynamical solution is unstable at fixed prey density, stable behavior cycles occur where neither forager nor prey keep a fixed behavior.