Neighbor intervention: a game-theoretic model

It has long been argued that a resident may benefit from helping its neighbor defend a territory against a challenger to avoid renegotiating its boundaries with a new and potentially stronger individual. We quantify this theory by exploring games involving challengers, residents and potential allies. In a simplified discrete game with zero variation of fighting strength, helping neighbors is part of an evolutionarily stable strategy (ESS) only if fighting costs are low relative to those of renegotiation. However, if relative fighting costs are high then an interventional ESS remains possible with finite variation of strength. Under these conditions, neighbors may help residents fight off intruders, but only when the resident does not stand a reliable chance of winning alone. We show that neighbor intervention is more likely with low home advantage to occupying a territory, strengths combining synergistically or low probability that an ally will be usurped, amongst other factors. Our parameterized model readily explains occasional intervention in the Australian fiddler crab, including why the ally tended to be larger than both the assisted neighbor and the intruder. Reciprocity is not necessary for this type of cooperation to persist, but also it is by no means inevitable in territorial species.     

The influence of contests on optimal clutch size: a game-theoretic model

We develop a game-theoretic model to predict the effect of size-dependent contest outcomes on optimal-clutch-size decisions. We consider the case where larger individuals develop from smaller clutches and, as adults, are advantaged in competition for limiting resources. The relationship between fitness and size thus depends on the sizes of other members of the population. We show that clutch-size optima are decreased by body-size-dependent contest outcomes, with larger effects when body size is most affected by clutch size, when prior resource ownership has less influence on contest outcome and when contests occur more frequently. We also show the existence of polymorphisms in clutch-size optima and that clutch-size driven changes in population density can, via an effect on the probability of host finding, further influence optimal clutch size. Our model is formulated to match the life history of a parasitoid wasp, in which clutch size affects offspring size and females engage in direct contests for host ownership, which larger females tend to win; we confirm that female-female competition is likely to influence clutch size in this species. However, the model is also relevant to clutch size in other taxa and supports recent suggestions concerning reproductive decisions in great tits.     

Altered aggression in different sized groups of crayfish supports a dynamic social behaviour model

Living in groups with conspecifics can increase an animal's fitness in the wild. A social environment may also be imposed by commercial farming industries. One important measure of competition and group dynamics is the level of aggressive interaction. This can also influence the level of damage or injury in cultured populations, a commercial issue at point of sale. There is considerable research into this issue in commercial species such as pigs, cattle and chickens but less is known about aquatic communal species such as decapod crustaceans. Here we manipulated group size in the freshwater crayfish Cherax destructor, a species that forms social groups in the wild and one that is also commercially farmed. Aggressive behaviour was scored during 1 h of observation in replicates of groups of 4, 16 and 36 animals to analyse 11 variables of fight dynamics that ensued. The number of fights per crayfish (4.0 ± 0.8 to 1.9 ± 0.2 fights, P = 0.017) and the time each crayfish was involved in a fight (113.9 ± 32.6 to 21.6 ± 2.6 s, P = 0.011) decreased as group size increased. Conversely, the number of failed tailflips elicited per crayfish increased from 0 to 0.08 ± 0.03 tailflips in the largest groups (P = 0.011). Together, the data suggest that despite C. destructor's different biology and habitat, compared to prior work that manipulates group size, the crayfish adjusts its fighting strategy when social circumstances change. Theory has proposed aggressive behaviour could change in groups of animals and our data indicates that this applies more broadly across species and more dynamically than previously demonstrated.     

Coalition formation: a game-theoretic analysis

There are many examples of individuals forming coalitions toobtain or protect a valuable resource. We present an analyticalmodel of coalition formation in which individuals seek alliancesif they judge themselves too weak to secure the resource alone.We allow coalition seeking to carry an investment cost () andlet contest outcomes depend probabilistically on the relativefighting strengths of contesting parties, with effective coalitionstrength directly proportional to combined partner strength.We identify the evolutionarily stable strength thresholds, belowwhich individuals within triads should seek a coalition. Weshow that if exceeds a critical value, then unilateral fightingover resources is an evolutionarily stable strategy (ESS). Universal(3-way) coalitions are also an ESS outcome if is less thana second critical value. Both of these extreme solutions areless likely to arise, the greater the variance in fighting strengthsand the greater the benefit from dominating opponents. Our analysisalso identifies intermediate solutions in which only the weakerindividuals seek coalitions: only then can a true coalition(2 vs. 1) form. We characterize these ESSs and show that truecoalitions are more likely to arise when the effective strengthof a coalition is less than the sum of its individual strengths(antergy). Alliances in primates are characterized by antergy,high reliability of strength as a predictor of contest outcome,and high variability in strengths. These are precisely the conditionsin which in our model most favors true coalition formation.     

A dynamic game-theoretic model of parental care

We present a model in which members of a mated pair decide whether to care for their offspring or desert them. There is a breeding season of finite length during which it is possible to produce and raise several batches of offspring. On deserting its offspring, an individual can search for a new mate. The probability of finding a mate depends on the number of individuals of each sex that are searching, which in turn depends upon the previous care and desertion decisions of all population members. We find the evolutionarily stable pattern of care over the breeding season. The feedback between behaviour and mating opportunity can result in a pattern of stable oscillations between different forms of care over the breeding season. Oscillations can also arise because the best thing for an individual to do at a particular time in the season depends on future behaviour of all population members. In the baseline model, a pair splits up after a breeding attempt, even if they both care for the offspring. In a version of the model in which a pair stays together if they both care, the feedback between behaviour and mating opportunity can lead to more than one evolutionarily stable form of care.     

The evolution of anisogamy: a game-theoretic approach

A popular theory has proposed that anisogamy originated through disruptive selection acting on an ancestral isogamous population, though recent work has emphasized the importance of other factors in its evolution. We re-examine the disruptive selection theory, starting from an isogamous population with two mating types and taking into account the functional relationship, g(m), between the fitness of a gamete and its size, m, as well as the relationship, f(S), between the fitness of a zygote and its size, S. Evolutionary game theory is used to determine the existence and continuous stability of isogamous and anisogamous strategies for the two mating types under various models for the two functions g(m) and f(S). In the ancestral unicellular state, these two functions are likely to have been similar; this leads to isogamy whether they are sigmoidal or concave, though in the latter case allowance must be made for a minimal gamete size. The development of multicellularity may leave g(m) relatively unchanged while f(S) moves to the right, leading to the evolution of anisogamy. Thus, the disruptive selection theory provides a powerful explanation of the origin of anisogamy, though other selective forces may have been involved in the subsequent specialization of micro- and macrogametes.     

On the evolution of pure winner and loser effects: A game-theoretic model

The persistence of linear dominance hierarchies is often attributed to higher probabilities of a win after a win or a loss after a loss in agonistic interactions, yet there has been no theory on the evolution of such prior-experience effects. Here an analytic model, based on the idea that contests are determined by subjective perceptions of resource-holding potential (RHP) which animals may revise in the light of experience, demonstrates that winner and loser effects can evolve through round-robin competition among triads of animals drawn randomly from their population, and that the probability of a hierarchy increases with the strength of the combined effect. The effects are pure, in the sense that a contestant observes neither its own RHP nor its opponent’s RHP or RHP perception or win—loss record; and so the strength of an effect is unmodified by the RHPs of particular individuals, but depends on the distribution of RHP among the population at large. The greater the difference between an individual’s and its opponent’s RHP perception, the more likely it is to win a contest; however, if it overestimates its RHP, then the cost of fighting increases with the overestimate. A winner or loser effect exists only if the fitness gain of the beta individual in a hierarchy, relative to that of the alpha, is less than 0.5. Then a loser effect can exist alone, or it can coexist with a winner effect; however, there cannot exist a winner effect without a loser effect.     

A game-theoretic model of kleptoparasitic behavior in polymorphic populations

Kleptoparasitism, the stealing of food by one animal from another, is a widespread biological phenomenon. In this paper we build upon earlier models to investigate a population of conspecifics involved in foraging and, potentially, kleptoparasitism. We assume that the population is composed of four types of individuals, according to their strategic choices when faced with an opportunity to steal and to resist an attack. The fitness of each type of individual depends upon various natural parameters, for example food density, the handling time of a food item and the probability of mounting a successful attack against resistance, as well as the choices that they make. We find the evolutionarily stable strategies (ESSs) for all parameter combinations and show that there are six possible ESSs, four pure and two mixtures of two strategies, that can occur. We show that there is always at least one ESS, and sometimes two or three. We further investigate the influence of the different parameters on when each type of solution occurs.     

The expected extinction time of a population within a system of interacting biological populations

The quasi-stationary distribution of a population within a system of interacting populations is approximated by a stochastic logistic process. The parameters of this process can be expressed in the parameters of the full system. Using the diffusion approximation, an expression for the expected extinction time is derived from this logistic process. Since the expected extinction time is expressed in the parameters of the full system, the effect of these parameters on the extinction risk can be easily evaluated, which may be of use for studies in ecology, conservation biology and epidemiology. The outcome is compared with simulation results for the case of a prey-predator system.     

Kin competition within groups: the offspring depreciation hypothesis

Where relatives compete for the same resources (kin competition) and each obtains an equal share, this can favour the evolution of elevated dispersal rates, such that most resource competition is among non-relatives. We show that this effect evaporates as among-sibling dominance increases to the point where the allocation of resources is maximally unequal. We restore a kin-competition effect on emigration rates from dominance-ranked family groups by showing that where siblings form queues to inherit the breeding positions, the length of the queue affects the fitness of all individuals by depreciating the rank of subsequent offspring. Incorporating this 'offspring depreciation' effect decreases optimal queue lengths, increases dispersal rates and explains the otherwise paradoxical use of sinks by cooperatively breeding birds in stable environments. The offspring depreciation effect also favours the evolution of small, but consistent, clutch sizes and high reproductive skew, but constrains the evolution of alloparenting.     

Depletion of SecDF-YajC causes a decrease in the level of SecG: implication for their functional interaction

SecA and an apparatus comprising SecYEG and SecDF-YajC complexes catalyze protein translocation across the Escherichia coli membrane. SecDF-YajC and SecG facilitate membrane insertion of SecA, which is the driving force for protein translocation. Here we report that SecDF-YajC depletion together with SecG depletion nearly completely inhibits protein translocation both in vivo and in vitro, although SecDF-YajC had been thought to be unnecessary for in vitro translocation. The level of SecG in membranes decreased to about half upon SecDF-YajC depletion and recovered to a normal level when SecDF-YajC was expressed. SecDF-YajC inhibited disulfide bond formation between two SecG molecules possessing a single cysteine residue. These results suggest functional interaction between SecDF-YajC and SecG.     

The ideal free distribution: a review and synthesis of the game-theoretic perspective

The Ideal Free Distribution (IFD), introduced by Fretwell and Lucas in [Fretwell, D.S., Lucas, H.L., 1970. On territorial behavior and other factors influencing habitat distribution in birds. Acta Biotheoretica 19, 16-32] to predict how a single species will distribute itself among several patches, is often cited as an example of an evolutionarily stable strategy (ESS). By defining the strategies and payoffs for habitat selection, this article puts the IFD concept in a more general game-theoretic setting of the “habitat selection game”. Within this game-theoretic framework, the article focuses on recent progress in the following directions: (1) studying evolutionarily stable dispersal rates and corresponding dispersal dynamics; (2) extending the concept when population numbers are not fixed but undergo population dynamics; (3) generalizing the IFD to multiple species.For a single species, the article briefly reviews existing results. It also develops a new perspective for Parker’s matching principle, showing that this can be viewed as the IFD of the habitat selection game that models consumer behavior in several resource patches and analyzing complications involved when the model includes resource dynamics as well. For two species, the article first demonstrates that the connection between IFD and ESS is now more delicate by pointing out pitfalls that arise when applying several existing game-theoretic approaches to these habitat selection games. However, by providing a new detailed analysis of dispersal dynamics for predator-prey or competitive interactions in two habitats, it also pinpoints one approach that shows much promise in this general setting, the so-called “two-species ESS”. The consequences of this concept are shown to be related to recent studies of population dynamics combined with individual dispersal and are explored for more species or more patches.     

Mathematical modeling of the course and prognosis of factitious disorders: a game-theoretic approach

A mathematical model using simple concepts of repeated games is proposed to model the course and prognosis of factitious disorders. Although simple, the model seems capable of explaining the yet unknown mechanisms underlying the variable course of factitious disorders. One of the notable results of this study is the significant effect of involved physicians in the treatment process on the course of the disease. Particularly, the doctor's error rate in realizing whether the symptoms are real or factitious can considerably affect the course of the disease. This is the first paper to apply a mathematical model to factitious disorders.     

Infrequent translation of a nonsense codon is sufficient to decrease mRNA level

In many organisms nonsense mutations decrease the level of mRNA. In the case of mammalian cells, it is still controversial whether translation is required for this nonsense-mediated RNA decrease (NMD). Although previous analyzes have shown that conditions that impede translation termination at nonsense codons also prevent NMD, the residual level of termination was unknown in these experiments. Moreover, the conditions used to impede termination might also have interfered with NMD in other ways. Because of these uncertainties, we have tested the effects of limiting translation of a nonsense codon in a different way, using two mutations in the immunoglobulin mu heavy chain gene. For this purpose we exploited an exceptional nonsense mutation at codon 3, which efficiently terminates translation but nonetheless maintains a high level of mu mRNA. We have shown 1) that translation of Ter462 in the double mutant occurs at only approximately 4% the normal frequency, and 2) that Ter462 in cis with Ter3 can induce NMD. That is, translation of Ter462 at this low (4%) frequency is sufficient to induce NMD.     

Deficiency of tenascin-X causes a decrease in the level of expression of type VI collagen

Tenascin-X (TNX) is an extracellular matrix glycoprotein. We previously demonstrated that TNX-null fibroblasts exhibit decreased cell-matrix and cell-cell adhesion. In this study, we used a differential display technique to determine the genes involved in this process. Differential display analysis of wild-type and TNX-null fibroblasts revealed that mRNA expression level of type VI collagen alpha3 is predominantly decreased in TNX-null fibroblasts. Expression levels of mRNAs of other subunits of type VI collagen, alpha2 and alpha3 chains, were also remarkably decreased in TNX-null fibroblasts. The protein level of alpha3 chain of type VI collagen was also reduced in TNX-null fibroblasts. However, the organization of type VI collagen in the extracellular matrix of TNX-null fibroblasts was similar to that of wild-type fibroblasts. Transient expression of TNX in Balb3T3 cells caused an increase in the level of mRNA of type VI collagen compared with that in vector control and increased the promoter activity of type VI collagen alpha1 subunit gene. In addition, the expression levels of type I collagen and other collagen fibril-associated molecules such as type XII and type XIV collagens, decorin, lumican and fibromodulin in wild-type and TNX-null fibroblasts were compared. It was found that the mRNA expression levels of type I collagen and collagen fibril-associated molecules other than decorin were decreased and that the expression level of decorin was increased in TNX-null fibroblasts. The results suggest the possibility that TNX mediates not only cell-cell and cell-matrix interactions but also fibrillogenesis via collagen fibril-associated molecules.     

Photosynthesis-irradiance response at physiological level: a mechanistic model

A mechanistic model is developed to present the photosynthetic response of phytoplankton to irradiance at the physiological level. The model is operated on photosynthetic units (PSU), and each PSU is assumed to have two states: reactive and activated. Light absorption that drives a reactive PSU into the activated state results from the effective absorption of the PSU. Transitions between the two states are asymmetrical in rate. A PSU in the reactive state becomes activated much faster than it recovers from the activated state to the reactive one. The turnover time for an activated PSU to transit into the reactive one is defined by the turnover time of the electron transport chain. The present model yields a photosynthesis-irradiance curve (PE-curve) in a hyperbola, which is described by three physiological parameters: effective cross-section (sigmaPSII), turnover time of electron transport chain (tau) and number of PSUs (N). The PE-curve has an initial slope of sigmaPSII x N, a half-saturated irradiance of 1/(tau sigmaPSII), and a maximal photosynthetic rate of N/tau at the saturated irradiance. The PE-curve from the present model is comparable to the empirical function based on the target theory described by the Poisson distribution.     

Results of pregnancies characterized by a decrease in the level of alpha-fetoprotein in the maternal blood

Pregnancy outcome was followed in 123 women showing maternal serum alpha-fetoprotein, less than or equal to 0.50 MOM. In 28 cases AFP was secondarily considered as normal either after ultrasonography and correction of gestation age or after a second sample normal result. In 95 cases AFP level was confirmed lowered; perinatal outcome was normal in 70 cases and abnormal in 25. Among these 25 cases, 3 autosomal trisomies occurred, 2 trisomies 18 and 1 trisomy 21; in the 22 other cases, we observed antepartum risk factors (10 cases with impending premature labor or premature labor, 9 cases with chronic hypertension, 2 cases with Ag HBs hepatitis and 1 case with diabetes).     

EEG correlates of aggression and anxiety in a social interaction model

Aggressiveness- and anxiety-related behavioral and oscillatory patterns were investigated in 49 18-30 year old subjects during virtual social interactions. The subjects were presented with pictures of "angry", "happy", and "neutral" faces and had to choose one out of three options: "attack", "avoid", or "make friends". Sources of cortical EEG were localized with sLORETA software. Subjects with high aggressiveness chose attack more frequently and this behavior was accompanied by a stronger induced delta and theta synchronization in the right orbitofrontal cortex. In subjects with high anxiety, delta and theta responses were stronger induced in the right temporal cortex during their more frequent avoidance behavior. Thus, both in anxious and in aggressive subjects, typical behavior was accompanied by increased induced low-frequency synchronization whose localization implies that it is associated with motivational and emotional processes.     

The expected probabilities of identity for the circular stepping-stone model

The expected values of the probabilities of identity by descent are derived for the circular stepping-stone model. The results are more easily interpreted than those derived previously.