Resistance is useless?—Extensions to the game theory of kleptoparasitism

We extend the game theoretic model of kleptoparasitism introduced by Broom and Ruxton (1998, Behav. Ecol. 9, 397–403) in two ways: we allow for asymmetric contests, where the probability α of the challenger winning can take any value from 0 to 1; and we allow the handler to choose not to resist the challenge, but to immediately concede and relinquish its food to the challenger. We find, in general, three possible evolutionarily stable strategies—challenge-and-resist (Hawk), challenge-but-do-not-resist (Marauder) and do-not-challenge-but-resist (Retaliator). When α = 1/2, we find that Hawk and Marauder are the only ESS’s, in contrast to the result of the original model; we also find an overlap region, in parameter space, where two different ESS’s are possible, depending on initial conditions. For general α, we see that all three ESS are possible, depending on different values of the environmental parameters; however, as the average time of a contest over food becomes long, then the Marauder strategy becomes more and more prevalent. The model makes a potentially significant prediction about animal behaviour in the area of kleptoparasitism, that a searcher, when it meets a handler, will only decline to attack that handler when α < 1/2 i.e.when the defender is more likely to win. One possible converse of this statement, that a handler whose probability of success is greater than 1/2 should always resist a challenge, is not true.     

Mathematical analysis of a model describing evolution of an asexual population in a changing environment

We investigate a mathematical model for an asexual population with non-overlapping (discrete) generations, that exists in a changing environment. Sexual populations are also briefly discussed at the end of the paper. It is assumed that selection occurs on the value of a single polygenic trait, which is controlled by a finite number of loci with discrete-effect alleles. The environmental change results in a moving fitness optimum, causing the trait to be subject to a combination of stabilising and directional selection.This model is different from that investigated by Waxman and Peck [Genetics 153 (1999) 1041] where overlapping generations and continuous effect alleles were considered. In this paper, we consider non-overlapping generations and discrete effect alleles. However in [Genetics 153 (1999) 1041] and the present work, there is the same pattern of environmental change, namely a constant rate of change of the optimum.From [Genetics 153 (1999) 1041], no rigorous theoretical conclusion can be drawn about the form of the solutions as t grows large. Numerical work carried out in [Genetics 153 (1999) 1041] suggests that the solution is a lagged travelling wave solution, but no mathematical proof exists for the continuous model. Only partial results, regarding existence of travelling wave solutions and perturbed solutions, have been established (see [Nonlin. Anal. 53 (2003) 683; An integral equation describing an asexual population in a changing environment, Preprint]). For the discrete case of this paper, under the assumption that the ratio between the unit of genotypic value and the speed of environment change is a rational number, we are able to give rigorous proof of the following conclusion: the population follows the environmental change with a small lag behind, moreover, the lag is represented using a calculable quantity.     

Modelling dominance hierarchy formation as a multi-player game

Animals who live in groups need to divide available resources amongst themselves. This is often achieved by means of a dominance hierarchy, where dominant individuals obtain a larger share of the resources than subordinate individuals. This paper introduces a model of dominance hierarchy formation using a multi-player extension of the classical Hawk-Dove game. Animals play non-independent pairwise games in a Swiss tournament which pairs opponents against those which have performed equally well in the conflict so far, for a fixed number of rounds. Resources are divided according to the number of contests won. The model, and its emergent properties, are discussed in the context of experimental observations.     

Evolution in knockout conflicts: The fixed strategy case

A group of individuals resolve their disputes by a knockout tournament. In each round of the tournament, the remaining contestants form pairs which compete, the winners progressing to the next round and the losers being eliminated. The payoff received depends upon how far the player has progressed and a cost is incurred only when it is defeated. We only consider strategies in which individuals are constrained to adopt a fixed play throughout the successive rounds. The case where individuals can vary their choice of behaviour from round to round will be treated elsewhere. The complexity of the system is investigated and illustrated both by special cases and numerical examples. M. Broom is also a member of the Centre for the Study of Evolution at the University of Sussex.     

Evolutionarily stable stealing: game theory applied to kleptoparasitism

We present an individual-based model of a group of foraginganimals. Individuals can obtain food either by discovering itthemselves or by stealing it from others (kleptoparasitism).Given that challenging another individual for a discovered fooditem costs time (which could otherwise be spent searching foran undiscovered item), attempting to steal from another maynot always be efficient We show that there is generally a uniquestrategy that maximizes uptake rate—always or never challengingothers. For any combination of parameter values, we can identifywhich strategy is appropraite. As a corollary to this, we predictthat small changes in ecolgical conditions can, under some circumstances,cause a dramatic change in the aggressive behavior of individuals.Further, we investigate situations where searching for undiscoveredfood and searching for potential opportunities for stealingare mutually exclusive activities (i.e., success at one canonly be improved at the expense of the other). Using game theory,we are able to find the evolutionarily stable strategy for investmentin these two activities in terms of the ecological parametersof the model.     

Exercise-induced suppression of acylated ghrelin in humans.

Ghrelin is an orexigenic hormone secreted from endocrine cells in the stomach and other tissues. Acylation of ghrelin is essential for appetite regulation. Vigorous exercise induces appetite suppression, but this does not appear to be related to suppressed concentrations of total ghrelin. This study examined the effect of exercise and feeding on plasma acylated ghrelin and appetite. Nine male subjects aged 19-25 yr participated in two, 9-h trials (exercise and control) in a random crossover design. Trials began at 0800 in the morning after an overnight fast. In the exercise trial, subjects ran for 60 min at 72% of maximum oxygen uptake between 0800 and 0900. After this, they rested for 8 h and consumed a test meal at 1100. In the control trial, subjects rested for 9 h and consumed a test meal at 1100. Area under the curve values for plasma acylated ghrelin concentration (assessed from venous blood samples) were lower over the first 3 h and the full 9 h of the exercise trial compared with the control trial: 317+/-135 vs. 510+/-186 pg.ml(-1).3 h and 917+/-342 vs. 1,401+/-521 pg.ml(-1).9 h (means+/-SE) respectively (P<0.05). Area under the curve values for hunger (assessed using a visual scale) were lower over the first 3 h of the exercise trial compared with the control trial (P=0.013). These findings demonstrate that plasma acylated ghrelin concentration and hunger are suppressed during running.     

Selective signalling by cuttlefish to predators

Rather than simply escaping, prey animals often attempt to deter an attack by signalling to an approaching predator, but this is a risky strategy if it allows time for the predator to draw closer (especially when the signal is a bluff). Because prey are vulnerable to multiple predators, the hunting techniques of which vary widely, it could well be beneficial for a prey animal to discriminate predators and to signal only to those that are likely to be deterred. Higher vertebrates make alarm calls that can identify the type of predator to the signaller's conspecifics, and a recent study shows that squirrels direct an infrared deterrent signal specifically at infrared-sensitive pit-vipers and not at other snakes. We show here that na?ve juvenile cuttlefish (Sepia officinalis L.) use a visual signal selectively during encounters with different predatory species. We analysed sequences of defensive behaviours produced by cuttlefish, to control for effects of relative threat level (or 'response urgency'). This showed that a high contrast 'eyespot' signal, known as the deimatic display, was used before flight against visually oriented teleost fish, but not crabs and dogfish, which are chemosensory predators.     

Rewired Metabolism in Drug-resistant Leukemia Cells: A METABOLIC SWITCH HALLMARKED BY REDUCED DEPENDENCE ON EXOGENOUS GLUTAMINE*

Cancer cells that escape induction therapy are a major cause of relapse. Understanding metabolic alterations associated with drug resistance opens up unexplored opportunities for the development of new therapeutic strategies. Here, we applied a broad spectrum of technologies including RNA sequencing, global untargeted metabolomics, and stable isotope labeling mass spectrometry to identify metabolic changes in P-glycoprotein overexpressing T-cell acute lymphoblastic leukemia (ALL) cells, which escaped a therapeutically relevant daunorubicin treatment. We show that compared with sensitive ALL cells, resistant leukemia cells possess a fundamentally rewired central metabolism characterized by reduced dependence on glutamine despite a lack of expression of glutamate-ammonia ligase (GLUL), a higher demand for glucose and an altered rate of fatty acid β-oxidation, accompanied by a decreased pantothenic acid uptake capacity. We experimentally validate our findings by selectively targeting components of this metabolic switch, using approved drugs and starvation approaches followed by cell viability analyses in both the ALL cells and in an acute myeloid leukemia (AML) sensitive/resistant cell line pair. We demonstrate how comparative metabolomics and RNA expression profiling of drug-sensitive and -resistant cells expose targetable metabolic changes and potential resistance markers. Our results show that drug resistance is associated with significant metabolic costs in cancer cells, which could be exploited using new therapeutic strategies.     

Optimal investment across different aspects of anti-predator defences

We present a simple model of investment across a suite of different anti-predatory defences. Defences can incur an initial construction cost and and/or may be costly each time they are utilised. Our aim is to use a simple, but general, mathematical model to explore when prey that face a single predatory threat where each attack is of the same nature should invest only in a single defence, and when they should spread their investment across more than one defence. This should help to explain the observed variety of defences that a single prey individual may employ during repeated attacks of a similar nature or even at different stages during one attack. Previous verbal reasoning suggested that prey should specialise in investment in defences that can be utilised early in the predation sequence. Our quantitative model predicts that (depending of the relatively properties of different defences), there may be concentrated investment in early acting, or in late-acting defences, or a spread of investment across both defence types. This variety of predictions is in agreement with the variation in defences shown by natural organisms subjected to repeated predatory attack.