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Biomedical subjects

A I Houston

Publications and source records attributed to A I Houston.

At least 19 recordsLinked to original sources

A dynamic model of hypothermia as an adaptive response by small birds to winter conditions.

We present a dynamic programming model which is used to investigate hypothermia as an adaptive response by small passerine birds in winter. The model predicts that there is a threshold function of reserves during the night, below which it is optimal to enter hypothermia, and above which it is optimal to rest. This threshold function decreases during the night, with a particularly sharp drop at the end of the night, representing the time and energy costs associated with returning to normal body temperature. The results of the model emphasise the trade-off between energy and predation, not just between foraging options, but also between foraging during the day and entering hypothermia at night. The value of being able to use hypothermia represents not just energy savings, but also reduced predation risk due to changes in the optimal foraging strategy. Conditions which give a high value of hypothermia are short photoperiod, variable food supply, low temperatures, poor and scarce food supplies.

Adaptation, Physiological↗

A theoretical investigation into the direct and indirect effects of state on the risk of predation.

As well as there being a direct physical effect of the state (for example fat reserves, or size) of an animal on the risk of being caught by a predator, state also has an effect on predation risk indirectly through changes in behaviour. We present a mathematical model which looks at these two components of the effect of state on predation risk. We focus on two different models, (i) where the animal must achieve a fixed state and its fitness depends on the time at which this state is reached and (ii) where the animal must survive until a fixed time and its fitness depends on its final state. We investigate conditions under which the indirect effect of increased state is to increase or decrease predation risk, and give some numerical illustrations. Under certain conditions in the fixed-state model, the indirect effect of state is to increase predation risk, whereas under certain conditions in the fixed-time model the indirect effect of state is to decrease predation risk. We discuss the implications of our results for empirical investigations into the effect of state on predation risk.

Animals↗

The strength of selection in the context of migration speed.

I use a model of avian migration based on maximization of overall migration speed to compare the strength of selection acting on foraging performance and flight speed. Let the optimal foraging behaviour be u* and the optimal flight speed be v*. It is shown that at this optimum, the ratio of the strength of selection on foraging to the strength of selection on flight speed is theta = -(u*2Pgamma"/v*2gammaP"), where gamma is the rate of energy expenditure during flight and P is the rate at which energy is gained during foraging. The dimensionless ratio P/gamma is the ratio of time spent building up fuel to time spent flying which A. Hedenström and T. Alerstam showed was much greater than unity. Although theta depends on this ratio, it also depends on the curvatures of the functions, as represented by gamma" and P". I use this simple example to make some general points about the strength of selection.

Animals↗

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.

Animals↗

Prey processing in central place foragers.

The importance of prey processing as an integral part of foraging behaviour has long been acknowledged, but little theoretical consideration has been given to the optimization of the processing behaviour itself. Processing renders food down to ingestible, palatable portions, and also removes non-essential mass thus reducing transport costs. Here, several models of processing are developed for a central place forager. When the forager has to make a simple choice between processing the prey and not, a critical distance from the central place can be calculated, beyond which it is optimal to process prey. If the forager also decides on how much of the prey to remove, the optimal amount to be removed can also be calculated. Imposing a ceiling on overall metabolic expenditure is shown to reduce the distances at which processing becomes the optimal strategy. The models are tested using parameters derived for a provisioning merlin, Falco columbarius, and alternative explanations as to why observed behaviours should differ from the optimal behaviour predicted are discussed.

Animals↗

Incorporating rules for responding into evolutionary games.

Evolutionary game theory is concerned with the evolutionarily stable outcomes of the process of natural selection. The theory is especially relevant when the fitness of an organism depends on the behaviour of other members of its population. Here we focus on the interaction between two organisms that have a conflict of interest. The standard approach to such two-player games is to assume that each player chooses a single action and that the evolutionarily stable action of each player is the best given the action of its opponent. We argue that, instead, most two-player games should be modelled as involving a series of interactions in which opponents negotiate the final outcome. Thus we should be concerned with evolutionarily stable negotiation rules rather than evolutionarily stable actions. The evolutionarily stable negotiation rule of each player is the best rule given the rule of its opponent. As we show, the action chosen as a result of the negotiation is not the best action given the action of the opponent. This conclusion necessitates a fundamental change in the way that evolutionary games are modelled.

Animals↗

A general technique for computing evolutionarily stable strategies based on errors in decision-making.

Realistic models of contests between animals will often involve a series of state-dependent decisions by the contestants. Computation of evolutionarily stable strategies for such state-dependent dynamic games are usually based on damped iterations of the best response map. Typically this map is discontinuous so that iterations may not converge and even if they do converge it may not be clear if the limiting strategy is a Nash equilibrium. We present a general computational technique based on errors in decision making that removes these computational difficulties. We show that the computational technique works for a simple example (the Hawk-Dove game) where an analytic solution is known, and prove general results about the technique for more complex games. It is also argued that there is biological justification for inclusion of the types of errors we have introduced.

Animals↗

State-dependent life histories.

Life-history theory is concerned with strategic decisions over an organism's lifetime. Evidence is accumulating about the way in which these decisions depend on the organism's physiological state and other components such as external circumstances. Phenotypic plasticity may be interpreted as an organism's response to its state. The quality of offspring may depend on the state and behaviour of the mother. Recent theoretical advances allow these and other state-dependent effects to be modelled within the same framework.

Aging↗

State-dependent life history evolution in Soay sheep: dynamic modelling of reproductive scheduling.

Adaptive decisions concerning the scheduling of reproduction in an animal's lifetime, including age at maturity and clutch or litter size, should depend on an animal's body condition or state. In this state-dependent case, we are concerned with the optimization of sequences of actions and so dynamic optimization techniques are appropriate. Here we show how stochastic dynamic programming can be used to study the reproductive strategies and population dynamics of natural populations, assuming optimal decisions. As examples we describe models based upon field data from an island population of Soay sheep on St. Kilda. This population shows persistent instability, with cycles culminating in high mortality every three or four years. We explore different assumptions about the extent to which Soay ewes use information about the population cycle in making adaptive decisions. We compare the observed distributions of strategies and population dynamics with model predictions; the results indicate that Soay ewes make optimal reproductive decisions given that they have no information about the population cycle. This study represents the first use of a dynamic optimization life history model of realistic complexity in the study of a field population. The techniques we use are potentially applicable to many other populations, and we discuss their extension to other species and other life history questions.

Acclimatization↗

Risk-sensitive foraging theory and operant psychology.

Hastjarjo, Silberberg, and Hursh (1990) have presented data on the foraging behavior of rats and discussed it in terms of risk-sensitive foraging theory. Because risk-sensitive foraging theory is comprised of several different models, it does not lead to general predictions about when an organism should prefer a foraging option with high variance to a foraging option with low variance. Any comparison of data with the predictions of the theory must be based on an appropriate model. I draw attention to various experiments that are potentially relevant to the results reported by Hastjarjo et al. and show how the time period over which the organism must survive can influence a model's predictions about risk sensitivity.

Animals↗

Genetic algorithms and evolution.

The genetic algorithm (GA) as developed by Holland (1975, Adaptation in Natural and Artificial Systems. Ann Arbor: University of Michigan Press) is an optimization technique based on natural selection. We use a modified version of this technique to investigate which aspects of natural selection make it an efficient search procedure. Our main modification to Holland's GA is the subdividing of the population into semi-isolated demes. We consider two examples. One is a fitness landscape with many local optima. The other is a model of singing in birds that has been previously analysed using dynamic programming. Both examples have epistatic interactions. In the first example we show that the GA can find the global optimum and that its success is improved by subdividing the population. In the second example we show that GAs can evolve to the optimal policy found by dynamic programming.

Algorithms↗

The value of fat reserves and the tradeoff between starvation and predation.

It is shown that in a range of models, the probability that a forager dies from starvation is, to a good approximation, an exponential function of energy reserves. Using a time and energy budget for a 19g passerine, we explore the consequences, in terms of starvation and predation, of various levels of energy reserves. It is shown that there exists an optimal level L* of reserves at which total mortality (starvation plus predation) is minimized. L* increases when the environment deteriorates as a result of a decrease in either temperature or mean gross gain or an increase in the mean search time. The effect of combined deteriorations is greater than the sum of their individual effects. At L*, the probability of predation is much higher than the probability of starvation. A simple analytic model suggests that this result will be fairly general, but also indicates conditions under which the result might not hold.

Adipose Tissue↗

Memory and the efficient use of information.

We consider the problem of how an animal's memory should be designed in order to cope with a stochastic and changing environment. In particular we consider the problem of forming the best estimate of an unknown and possibly changing environmental parameter. Under the simple model we consider, the effect of an observation is to update this estimate using a linear operator. Two models of a changing environment are analysed. For each model we show how estimates change as a function of time elapsed and observations taken. The effect of a regular sequence of observations is also considered, and it is shown that an exponential weighting of past observations is a sufficient statistic on which to base decisions. The weighting factors are different in the two model environments considered, but each is shown to be a function of the rate at which the environment is changing.

Animals↗

A general framework for understanding the effects of variability and interruptions on foraging behaviour.

A general framework for analysing the effects of variability and the effects of interruptions on foraging is presented. The animal is characterised by its level of energetic reserves, x. We consider behaviour over a period of time [O,T]. A terminal reward function R(x) determines the expected future reproductive success of an animal with reserves x at time T. For any state x at a time in the period, we give the animal a choice between various options and then constrain it to follow a background strategy. The best option is the one that maximizes expected future reproductive success. Using this framework, we show that sensitivity to variability in amount of energy gained is logically distinct from sensitivity to variability in the time at which food is obtained. We also show that incorporating interruptions results in both a preference for variability in time and a preference for a reward followed by a delay as opposed to the same delay before the reward.

Animals↗

Relative allocation on concurrent schedules can depend on schedule parameters when behavioral parameters are constant.

We develop a simple model of switching between the initial links of a concurrent-chain procedure. Behavior is determined by four parameters mu(1), mu(2), q(1), and q(2). The first two are the basic rates of switching from Schedule 1 and Schedule 2, respectively. The second two are the probabilities of leaving Schedule 1 and Schedule 2 after the corresponding terminal link has been completed. We show that for fixed values of these four parameters, the relative allocation on the initial links may change as a result of changes in initial-link schedules. The effect can be quite large if the switching rates are low. An implication is that relative allocation is not necessarily a good measure of behavior.

Journal Article↗

The maximization of overall reinforcement rate on concurrent chains.

We model behavioral allocation on concurrent chains in which the initial links are independent variable-interval schedules. We also quantify the relationship between behavior during the initial links and the probability of entering a terminal link. The behavior that maximizes overall reinforcement rate is then considered and compared with published experimental data. Although all the trends in the data are predicted by rate maximization, there are considerable deviations from the predictions of rate maximization when reward magnitudes are unequal. We argue from our results that optimal allocation on concurrent chains, and prey choice as used in the theory of optimal diets, are distinct concepts. We show that the maximization of overall rate can lead to apparent violations of stochastic transitivity.

Journal Article↗

Another look at the control of behaviour by internal and external factors.

Toates (1981) claims that internal and external factors interact in the control of behaviour. This leads him to criticise the framework developed by McFarland and Sibly (1975) on the grounds that they assume that causal factors are independent. This paper argues that "independence" as used by McFarland and Sibly does not preclude the possibility of interactions as envisaged by Toates. It is then argued that it may be possible to represent the apparent examples of interactions by non-interacting causal factors.

Animals↗

The case against Allen's generalization of the matching law.

Allen (1981) claims to have established the formal validity of a power-function generalization of the matching law. This paper argues that Allen's proof is not correct when schedules are presented in pairs and that his initial assumptions are too restrictive when all schedules are simultaneously available.

Journal Article↗