Group selection, individual selection, and the evolution of genetic drift.

In a subdivided population, genetic drift affects variation between groups, and thus it can have an important effect on the outcome of evolution (Wright, 1978). The rate of genetic drift is determined, in part, by the behaviour of population members. This paper presents three mathematical models in which behavioural traits that affect the rate of genetic drift are allowed to coevolve with traits that are under selection at the group and individual levels. The results show that if group selection is strong relative to individual selection, then behavioural traits that enhance the rate of genetic drift will tend to increase in frequency. The strength of this effect depends, in part, on the way in which vacant sites are colonized.     

The evolution of brain lateralization: a game-theoretical analysis of population structure

In recent years, it has become apparent that behavioural and brain lateralization at the population level is the rule rather than the exception among vertebrates. The study of these phenomena has so far been the province of neurology and neuropsychology. Here, we show how such research can be integrated with evolutionary biology to understand lateralization more fully. In particular, we address the fact that, within a species, left- and right-type individuals often occur in proportions different from one-half (e.g. hand use in humans). The traditional explanations offered for lateralization of brain function (that it may avoid unnecessary duplication of neural circuitry and reduce interference between functions) cannot account for this fact, because increased individual efficiency is unrelated to the alignment of lateralization at the population level. A further puzzle is that such an alignment may even be disadvantageous, as it makes individual behaviour more predictable to other organisms. Here, we show that alignment of the direction of behavioural asymmetries in a population can arise as an evolutionarily stable strategy when individual asymmetrical organisms must coordinate their behaviour with that of other asymmetrical organisms. Brain and behavioural lateralization, as we know it in humans and other vertebrates, may have evolved under basically 'social' selection pressures.     

Computational models of spatial updating in peri-saccadic perception

Perceptual phenomena that occur around the time of a saccade, such as peri-saccadic mislocalization or saccadic suppression of displacement, have often been linked to mechanisms of spatial stability. These phenomena are usually regarded as errors in processes of trans-saccadic spatial transformations and they provide important tools to study these processes. However, a true understanding of the underlying brain processes that participate in the preparation for a saccade and in the transfer of information across it requires a closer, more quantitative approach that links different perceptual phenomena with each other and with the functional requirements of ensuring spatial stability. We review a number of computational models of peri-saccadic spatial perception that provide steps in that direction. Although most models are concerned with only specific phenomena, some generalization and interconnection between them can be obtained from a comparison. Our analysis shows how different perceptual effects can coherently be brought together and linked back to neuronal mechanisms on the way to explaining vision across saccades.     

Quantitative descriptions of resource choice in ecological models

This article reviews the subject of resource choice by consumers. It is concerned with how such choice has been and should be represented in quantitative ecological models. This requires consideration of the dynamics of behavioral change and the fitness consequences of different resource intake rates. The topic is important because of the impact of choice on the functional response of the consumer to each of the resources it consumes. A variety of open questions related to choice are addressed. These include: the relationship between optimal diet choice and switching; the relationship between adaptive choice of two or resources and type-3 functional responses to a single resource; whether switching behavior requires choice and whether choice always results in switching behavior; why partial preferences are observed; whether choice between habitats is fundamentally different from choice within habitats; how between-individual variation in parameters related to resource use alters functional responses measured at the population level. The impacts of choice on stability are discussed briefly. The costs of increased resource use and the type of nutritional interactions between resources are particularly important determinants of adaptive resource choice, and are considered in some detail.     

New developments in solid-state fermentation: II. Rational approaches to the design, operation and scale-up of bioreactors

Over the last decade there has been a significant improvement in understanding how to design, operate and scale-up solid-state fermentation bioreactors. The key to these advances has been the application of mathematical modeling techniques to describe the biological and transport phenomena within the system. This review focuses on the advances in understanding that have come from this modeling work, and the insights it has given us into bioreactor design, operation and scale-up. It also highlights two promising bioreactor designs that have emerged over the last decade or so. For processes in which the substrate bed must remain static throughout the fermentation, the most promising design is the Zymotis design of ORSTOM at Montpellier, France, which involves closely spaced internal heat transfer plates within a packed-bed bioreactor. For those processes in which mixing can be tolerated, the stirred bioreactor developed at INRA, in Dijon, France, has been successfully demonstrated at scales of 1–25 t of substrate. Theoretical work suggests that mathematical models will be useful tools in the scale-up process, however, there are no reports that they have been used in the development of any current large-scale process. Rather, the models have been validated against data obtained from laboratory-scale bioreactors. There is an urgent need to test the accuracy and robustness of the models by applying them within real process development.     

Multiscale modeling and mechanics of filamentous actin cytoskeleton

The adaptive structure and functional changes of the actin cytoskeleton are induced by its mechanical behavior at various temporal and spatial scales. In particular, the mechanical behaviors at different scales play important roles in the mechanical functions of various cells, and these multiscale phenomena require clarification. To establish a milestone toward achieving multiscale modeling and simulation, this paper reviews mathematical analyses and simulation methods applied to the mechanics of the filamentous actin cytoskeleton. The actin cytoskeleton demonstrates characteristic behaviors at every temporal and spatial scale, and mathematical models and simulation methods can be applied to each level of actin cytoskeletal structure ranging from the molecular to the network level. This paper considers studies on mathematical models and simulation methods based on the molecular dynamics, coarse-graining, and continuum dynamics approaches. Every temporal and spatial scale of actin cytoskeletal structure is considered, and it is expected that discrete and continuum dynamics ranging from functional expression at the molecular level to macroscopic functional expression at the whole cell level will be developed and applied to multiscale modeling and simulation.     

Predicting the effect of disturbance on coastal birds

Assessments of whether disturbance is having a deleterious effect on populations have often measured behavioural responses to disturbance and assumed that populations with a larger behavioural response are more susceptible to disturbance. However, there is no guarantee that the behavioural response to disturbance is related to the population consequence, measured in terms of decreased reproduction or increased mortality. Individual-based models, consisting of fitness-maximizing individuals, are one means of linking the behavioural responses to disturbance to population consequences. This paper reviews how individual-based models have been used to predict the effect of disturbance on populations of shorebirds and wildfowl at several European sites, and shows how these models could be improved in the future by incorporating a range of alternative responses to disturbance.     

Individual differences in behavioural plasticities

Interest in individual differences in animal behavioural plasticities has surged in recent years, but research in this area has been hampered by semantic confusion as different investigators use the same terms (e.g. plasticity, flexibility, responsiveness) to refer to different phenomena. The first goal of this review is to suggest a framework for categorizing the many different types of behavioural plasticities, describe examples of each, and indicate why using reversibility as a criterion for categorizing behavioural plasticities is problematic. This framework is then used to address a number of timely questions about individual differences in behavioural plasticities. One set of questions concerns the experimental designs that can be used to study individual differences in various types of behavioural plasticities. Although within‐individual designs are the default option for empirical studies of many types of behavioural plasticities, in some situations (e.g. when experience at an early age affects the behaviour expressed at subsequent ages), ‘replicate individual’ designs can provide useful insights into individual differences in behavioural plasticities. To date, researchers using within‐individual and replicate individual designs have documented individual differences in all of the major categories of behavioural plasticities described herein. Another important question is whether and how different types of behavioural plasticities are related to one another. Currently there is empirical evidence that many behavioural plasticities [e.g. contextual plasticity, learning rates, IIV (intra‐individual variability), endogenous plasticities, ontogenetic plasticities) can themselves vary as a function of experiences earlier in life, that is, many types of behavioural plasticity are themselves developmentally plastic. These findings support the assumption that differences among individuals in prior experiences may contribute to individual differences in behavioural plasticities observed at a given age. Several authors have predicted correlations across individuals between different types of behavioural plasticities, i.e. that some individuals will be generally more plastic than others. However, empirical support for most of these predictions, including indirect evidence from studies of relationships between personality traits and plasticities, is currently sparse and equivocal. The final section of this review suggests how an appreciation of the similarities and differences between different types of behavioural plasticities may help theoreticians formulate testable models to explain the evolution of individual differences in behavioural plasticities and the evolutionary and ecological consequences of individual differences in behavioural plasticities.     

Partner choice creates fairness in humans

Many studies demonstrate that partner choice has played an important role in the evolution of human cooperation, but little work has tested its impact on the evolution of human fairness. In experiments involving divisions of money, people become either over-generous or over-selfish when they are in competition to be chosen as cooperative partners. Hence, it is difficult to see how partner choice could result in the evolution of fair, equal divisions. Here, we show that this puzzle can be solved if we consider the outside options on which partner choice operates. We conduct a behavioural experiment, run agent-based simulations and analyse a game-theoretic model to understand how outside options affect partner choice and fairness. All support the conclusion that partner choice leads to fairness only when individuals have equal outside options. We discuss how this condition has been met in our evolutionary history, and the implications of these findings for our understanding of other aspects of fairness less specific than preferences for equal divisions of resources.     

Modelling spatio-temporal interactions within the cell

Biological phenomena at the cellular level can be represented by various types of mathematical formulations. Such representations allow us to carry out numerical simulations that provide mechanistic insights into complex behaviours of biological systems and also generate hypotheses that can be experimentally tested. Currently, we are particularly interested in spatio-temporal representations of dynamic cellular phenomena and how such models can be used to understand biological specificity in functional responses. This review describes the capability and limitations of the approaches used to study spatio-temporal dynamics of cell signalling components.     

Simplified models for gas exchange in the human lungs

This paper presents a hierarchy of models with increasing complexity for gas exchange in the human lungs. The models span from a single compartment, inflexible lung to a single compartment, flexible lung with pulmonary gas exchange. It is shown how the models are related to well-known models in the literature. A long-term purpose of this work is to study nonlinear phenomena seen in the cardio-respiratory system (for example, synchronization between ventilation rate and heart rate, and Cheyne-Stokes respiration). The models developed in this paper can be regarded as the controlled system (plant) and provide a mathematical framework to link between "molecular-level", and "systems-level" models. It is shown how changes in molecular level affect the alveolar partial pressure. Two assumptions that have previously been made are re-examined: (1) the hidden assumption that the air flow through the mouth is equal to the rate of volume change in the lungs, and, (2) the assumption that the process of oxygen binding to hemoglobin is near equilibrium. Conditions under which these assumptions are valid are studied. All the parameters in the models, except two, are physiologically realistic. Numerical results are consistent with published experimental observations.     

What we talk about when we talk about access deficits

Semantic impairments have been divided into storage deficits, in which the semantic representations themselves are damaged, and access deficits, in which the representations are intact but access to them is impaired. The behavioural phenomena that have been associated with access deficits include sensitivity to cueing, sensitivity to presentation rate, performance inconsistency, negative serial position effects, sensitivity to number and strength of competitors, semantic blocking effects, disordered selection between strong and weak competitors, correlation between semantic deficits and executive function deficits and reduced word frequency effects. Four general accounts have been proposed for different subsets of these phenomena: abnormal refractoriness, too much activation, impaired competitive selection and deficits of semantic control. A combination of abnormal refractoriness and impaired competitive selection can account for most of the behavioural phenomena, but there remain several open questions. In particular, it remains unclear whether access deficits represent a single syndrome, a syndrome with multiple subtypes or a variable collection of phenomena, whether the underlying deficit is domain-general or domain-specific, whether it is owing to disorders of inhibition, activation or selection, and the nature of the connection (if any) between access phenomena in aphasia and in neurologically intact controls. Computational models offer a promising approach to answering these questions.     

Pressure oscillation delivery to the lung: Computer simulation of neonatal breathing parameters

Preterm newborn infants may develop respiratory distress syndrome (RDS) due to functional and structural immaturity. A lack of surfactant promotes collapse of alveolar regions and airways such that newborns with RDS are subject to increased inspiratory effort and non-homogeneous ventilation. Pressure oscillation has been incorporated into one form of RDS treatment; however, how far it reaches various parts of the lung is still questionable. Since in-vivo measurement is very difficult if not impossible, mathematical modeling may be used as one way of assessment. Whereas many models of the respiratory system have been developed for adults, the neonatal lung remains essentially ill-described in mathematical models. A mathematical model is developed, which represents the first few generations of the tracheo-bronchial tree and the 5 lobes that make up the premature ovine lung. The elements of the model are derived using the lumped parameter approach and formulated in Simulink? within the Matlab? environment. The respiratory parameters at the airway opening compare well with those measured from experiments. The model demonstrates the ability to predict pressures, flows and volumes in the alveolar regions of a premature ovine lung.     

Mechanisms of sperm competition: Linking physiology and behavioural ecology

Sperm competition has received a great deal of attention from behavioural ecologists because it is the ultimate form of male-male competition, and may also be important for female choice. It is becoming clear that the adaptive value of the behavioural strategies of males and females will not be fully understood until we have a better understanding of the physiological mechanisms that come into play after copulation. We now have enough information to compare the underlying mechanisms in birds and mammals and to relate these to the way in which sperm competition operates in both groups. To integrate this knowledge, the boundaries between behavioural ecology and physiology will have to be crossed in what promises to be a most fruitful enterprise.     

Designing mate choice experiments

The important role that mate choice plays in the lives of animals is matched by the large and active research field dedicated to studying it. Researchers work on a wide range of species and behaviours, and so the experimental approaches used to measure animal mate choice are highly variable. Importantly, these differences are often not purely cosmetic; they can strongly influence the measurement of choice, for example by varying the behaviour of animals during tests, the aspects of choice actually measured, and statistical power. Consideration of these effects are important when comparing results among studies using different types of test, or when using laboratory results to predict animal behaviour in natural populations. However, these effects have been underappreciated by the mate choice literature to date. I focus on five key experimental considerations that may influence choice: (i) should mating be allowed to occur, or should a proxy behavioural measure of preference be used instead? (ii) Should subjects be given a choice of options? (iii) Should each subject be tested more than once, either with the same or different stimuli? (iv) When given a choice, how many options should the subject choose between? (v) What form should the experimental stimuli take? I discuss the practical advantages and disadvantages of common experimental approaches, and how they may influence the measurement of mate choice in systematic ways. Different approaches often influence the ability of animals to perceive and compare stimuli presented during tests, or the perceived costs and benefits of being choosy. Given that variation in the design of mate choice experiments is likely unavoidable, I emphasise that there is no single ‘correct’ approach to measuring choice across species, although ecological relevance is crucial if the aim is to understand how choice acts in natural populations. I also highlight the need for quantitative estimates of the sizes of potentially important effects, without which we cannot make informed design decisions.     

Is more choice always desirable? Evidence and arguments from leks,food selection,and environmental enrichment

Recent studies on humans show that too much choice can make subjects less likely to choose any item. I consider general adaptive and non‐adaptive explanations of why such choice aversion, or its converse, might occur in animals. There are three questions: is more choice always preferred, does it ever lead to less consumption (or a lower probability of consumption), and may it result in worse items being selected? A preference for choice is one of the main explanations for lek formation and I draw attention to previously unrecognised parallels with models of human shopping behaviour. There is indeed evidence of female preference for larger leks, although much of the observational data are open to other interpretations. Unfortunately nobody has looked for choice aversion where it is most to be expected, in leks larger than normally occur. Evidence that too much choice of males confuses females is strongest in acoustically advertising frogs, but the widespread decrease of mating skew in larger leks might also have this explanation. A model reanalyses data on skew in black grouse Tetrao tetrix and suggests that considering only a random subset of a large lek may increase the chances of selecting the better males: larger leks are more likely to include better males, but these are less likely to be selected. These opposing effects may lead to an optimum lek size, but only with a sufficient decline in choice accuracy with size. With food choice, very few studies have avoided confounding choice with food quality, by manipulating only flavour. The widespread phenomena of stimulus‐specific satiety and novelty seeking imply that monotonous diets are aversive, but no studies test whether animals choose sites where they know food diversity to be greater. Operant experiments that demonstrate mild preferences for free choice concern choice about the means to get food rather than the food itself. In some insect species even moderate choice of diet can be deleterious, and studies on search images and the confusion effect may be evidence of this in vertebrates. Environmental enrichment of captive animals often relies on increasing the options available, but it need not be the choice itself that is beneficial. I consider briefly further areas in biology where choice preference or aversion are potentially important.     

Mathematical Models for Sleep-Wake Dynamics: Comparison of the Two-Process Model and a Mutual Inhibition Neuronal Model

Sleep is essential for the maintenance of the brain and the body, yet many features of sleep are poorly understood and mathematical models are an important tool for probing proposed biological mechanisms. The most well-known mathematical model of sleep regulation, the two-process model, models the sleep-wake cycle by two oscillators: a circadian oscillator and a homeostatic oscillator. An alternative, more recent, model considers the mutual inhibition of sleep promoting neurons and the ascending arousal system regulated by homeostatic and circadian processes. Here we show there are fundamental similarities between these two models. The implications are illustrated with two important sleep-wake phenomena. Firstly, we show that in the two-process model, transitions between different numbers of daily sleep episodes can be classified as grazing bifurcations. This provides the theoretical underpinning for numerical results showing that the sleep patterns of many mammals can be explained by the mutual inhibition model. Secondly, we show that when sleep deprivation disrupts the sleep-wake cycle, ostensibly different measures of sleepiness in the two models are closely related. The demonstration of the mathematical similarities of the two models is valuable because not only does it allow some features of the two-process model to be interpreted physiologically but it also means that knowledge gained from study of the two-process model can be used to inform understanding of the behaviour of the mutual inhibition model. This is important because the mutual inhibition model and its extensions are increasingly being used as a tool to understand a diverse range of sleep-wake phenomena such as the design of optimal shift-patterns, yet the values it uses for parameters associated with the circadian and homeostatic processes are very different from those that have been experimentally measured in the context of the two-process model.     

Behavioural social choice: a status report

Behavioural social choice has been proposed as a social choice parallel to seminal developments in other decision sciences, such as behavioural decision theory, behavioural economics, behavioural finance and behavioural game theory. Behavioural paradigms compare how rational actors should make certain types of decisions with how real decision makers behave empirically. We highlight that important theoretical predictions in social choice theory change dramatically under even minute violations of standard assumptions. Empirical data violate those critical assumptions. We argue that the nature of preference distributions in electorates is ultimately an empirical question, which social choice theory has often neglected. We also emphasize important insights for research on decision making by individuals. When researchers aggregate individual choice behaviour in laboratory experiments to report summary statistics, they are implicitly applying social choice rules. Thus, they should be aware of the potential for aggregation paradoxes. We hypothesize that such problems may substantially mar the conclusions of a number of (sometimes seminal) papers in behavioural decision research.