How host plant variability influences the advantages to learning: a theoretical model for oviposition behaviour in Lepidoptera

Learning can allow individuals to increase their fitness in particular environments. The advantage to learning depends on the predictability of the environment and the extent to which animals can adjust their behaviour. Earlier general models have investigated when environmental predictability might favour the evolution of learning in foraging animals. Here, we construct a theoretical model that predicts the advantages to learning using a specific biological example: oviposition in the Lepidoptera. Our model includes environmental and behavioural complexities relevant to host selection in these insects and tests whether the predictions of the general models still hold. Our results demonstrate how the advantage of learning is maximised when within-generation variability is minimised (the local environment consists mainly of a single host plant species) and between-generation variability is maximised (different host plant species are the most common in different generations). We discuss how our results: (a) can be applied to recent empirical work in different lepidopteran species and (b) predict an important role of learning in lepidopteran agricultural pests.     

Within‐generation bet hedging: a seductive explanation?

Bet hedging occurs when a single genotype shows a variety of phenotypes in the same environment, and each phenotype is successful only when the particular circumstances to which it is adapted occur. The time scale of between-generation bet hedging ensures that all individuals with a given phenotype suffer the same fate – circumstances such as drought exert homogenous pressure on all members of a population. Under within-generation bet hedging, however, individuals with the same phenotype are subject to heterogeneous selection pressure – predation, for example, will affect some individuals but not others. An important consequence of this difference is that conditions favoring the evolution of within-generation bet hedging are very restricted. While a single lineage may realize increased fitness via within-generation bet hedging, this fitness advantage varies inversely with population size and becomes vanishingly small at even modest population sizes. Although several reviews and analyses have highlighted the differences between these two types of bet hedging, confusion persists regarding their respective definitions and evolutionary justification. Bet hedging is a seductive explanation because most students of evolution are trained to focus on costs and benefits at the individual level, and tend to seek adaptive explanations for individual traits. Although this focus is often successful, it leads us astray in the case of within-generation bet hedging. Only by assessing the fitness effects of a trait in the context of whole populations can one accurately identify traits that can and cannot be favored by within-generation bet hedging.     

The selective advantage of reaction norms for environmental tolerance

A tolerance curve defines the dependence of a genotype's fitness on the state of an environmental gradient. It can be characterized by a mode (the genotype's optimal environment) and a width (the breadth of adaptation). It seems possible that one or both of these characters can be modified in an adaptive manner, at least partially, during development. Thus, we extend the theory of environmental tolerance to include reaction norms for the mode and the width of the tolerance curve. We demonstrate that the selective value of such reaction norms increases with increasing spatial heterogeneity and between-generation temporal variation in the environment and with decreasing within-generation temporal variation. Assuming that the maintenance of a high breadth of adaptation is costly, reaction, norms are shown to induce correlated selection for a reduction in this character. Nevertheless, regardless of the magnitude of the reaction norm, there is a nearly one to one relationship between the optimal breadth of adaptation and the within-generation temporal variation perceived by the organism. This suggests that empirical estimates of the breadth of adaptation may provide a useful index of this type of environmental variation from the organism's point of view.     

Molecular Variation and Possible Lines Of Evolution of Peptide and Protein Hormones

The widespread distribution of certain steroids and amino acidderivatives with hormonal properties is considered evidencein support of the dictum that "it is not the hormones that change,but rather the uses to which they are put." However, analysesof the distributions, biological activities, immunological cross-reactivities,and sequences of amino acids of five representative peptideand protein hormones or groups of hormones—lactogenichormone, growth hormone, the corticotropin-MSH-ß lipotropinfamily, insulin, and the neurohypophysial hormones—supporta concept of change and of molecular evolution of these polypeptidicmolecules. When analyzed in terms of the genetic code, the aminoacid interchanges which have been revealed by determinationof sequences of amino acids can, most often, be explained bysingle base mutations in the appropriate codons. In two instanceswhere two base mutations within a single codon are required,intermediate replacements of amino acid have been suggested;one of these would lead to a 2-ALA-ß MSH, and theother to a 4-PRO, 8-ILE oxytocin.     

Persistence of costly novel genes in the absence of positive selection

Many genetic changes that ultimately lead to adaptive evolution come with a short‐term cost expressed in terms of reduced survival and reproduction. In the absence of genetic drift, it is unclear how such costly mutations may persist. Here we experimentally demonstrate that parasites can promote the persistence of costly genetic variants. We employed a genetically engineered strain (GMMO) of the bacterium Pseudomonas fluorescens as a model of the acquisition of a new gene either through a major mutation or through horizontal transfer, and examined its persistence in different evolving communities comprising an ancestral strain and a lytic bacteriophage. Whereas competition resulted in the elimination of the GMMO, inclusion of the phage promoted GMMO persistence. We provide evidence for why this effect is due to the differential susceptibility of GMMO and ancestral bacteria to phage.     

Expanding networks: Signaling components in and a hypothesis for the evolution of metamorphosis

Metamorphosis is a substantial morphological transition between2 multicellular phases in an organism's life cycle, often markingthe passage from a prereproductive to a reproductive life stage.It generally involves major physiological changes and a shiftin habitat and feeding mode, and can be subdivided into an extendedphase of substantial morphological change and/or remodeling,and a shorter-term phase (for example, marine invertebrate "settlement,"insect "adult eclosion," mushroom fruiting body emergence) wherethe actual habitat shift occurs. Disparate metamorphic taxadiffer substantially with respect to when the habitat shiftoccurs relative to the timing of the major events of morphogeneticchange. I will present comparative evidence across a broad taxonomicscope suggesting that longer-term processes (morphogenetic changes)are generally hormonally regulated, whereas nitric oxide (NO)repressive signaling often controls the habitat shift itself.Furthermore, new evidence from echinoids (sea urchins, sanddollars) indicates a direct connection between hormonal andNO signaling during metamorphosis. I incorporate 2 hypothesesfor the evolution of metamorphosis—one involving heterochrony,the other involving phenotypic integration and evolutionarilystable configurations (ESCs)—into a network model formetamorphosis in echinoderms (sea urchins, starfish, and theirkin). Early indications are that this core regulatory networkcan be acted upon by natural selection to suit the diverse ecologicalneeds of disparate metamorphic organisms, resulting in evolutionaryexpansions and contractions in the core network. I briefly speculateon the ways that exposure to xenobiotic pollutants and othercompounds might influence successful settlement of juvenilesin the wild. Indeed, environmentally regulated life historytransitions—such as settlement, metamorphosis, and reproductivematuration—may be developmental periods that are especiallysensitive to such pollutants.     

Learning in a game context: strategy choice by some keeps learning from evolving in others

Behavioural decisions in a social context commonly have frequency-dependent outcomes and so require analysis using evolutionary game theory. Learning provides a mechanism for tracking changing conditions and it has frequently been predicted to supplant fixed behaviour in shifting environments; yet few studies have examined the evolution of learning specifically in a game-theoretic context. We present a model that examines the evolution of learning in a frequency-dependent context created by a producer–scrounger game, where producers search for their own resources and scroungers usurp the discoveries of producers. We ask whether a learning mutant that can optimize its use of producer and scrounger to local conditions can invade a population of non-learning individuals that play producer and scrounger with fixed probabilities. We find that learning provides an initial advantage but never evolves to fixation. Once a stable equilibrium is attained, the population is always made up of a majority of fixed players and a minority of learning individuals. This result is robust to variation in the initial proportion of fixed individuals, the rate of within- and between-generation environmental change, and population size. Such learning polymorphisms will manifest themselves in a wide range of contexts, providing an important element leading to behavioural syndromes.     

Rhizobial communities in symbiosis with legumes: genetic diversity,competition and interactions with host plants

The term ‘Rhizobium-legume symbiosis’ refers to numerous plant-bacterial interrelationships. Typically, from an evolutionary perspective, these symbioses can be considered as species-to-species interactions, however, such plant-bacterial symbiosis may also be viewed as a low-scale environmental interplay between individual plants and the local microbial population. Rhizobium-legume interactions are therefore highly important in terms of microbial diversity and environmental adaptation thereby shaping the evolution of plant-bacterial symbiotic systems. Herein, the mechanisms underlying and modulating the diversity of rhizobial populations are presented. The roles of several factors impacting successful persistence of strains in rhizobial populations are discussed, shedding light on the complexity of rhizobial-legume interactions.     

Structure of Desert Rodent Communities: A Critical Review of Questions and Approaches

Three attributes of communities—the number, relative abundances,and phenotypic attributes of coexisting species—togetherdefine their "structure," as the term has been used in the literature.Most ecologists have tried to uncover determinants of communitystructure by analyzing patterns of morphology or resource use,or by experimentally manipulating the species composition orenvironmental context of communities. Less often used is a mechanisticanalysis of processes operating at the level of individualsor populations. I discuss the logical basis for each of these approaches andillustrate their virtues and limitations with examples drawnfrom work on heteromyid rodents. While pattern analytic andexperimental approaches have provided an efficient means ofidentifying proximate determinants of species number and relativeabundance, a mechanistic approach holds more promise for answeringquestions about the ultimate determinants of phenotypic patternswithin communities     

Colony Specificity in the Colonial Tunicate Botryllus and the Origins of Vertebrate Immunity

SYNOPSIS. Colonies of the compound tunicate Botryllus show thecapacity for self—nonself discrimination by fusion betweenseparated pieces of the same colony and rejection between piecesof unrelated colonies. We have found that genes controllingthis colony specificity are similar to those which cause transplantrejection in the vertebrates. Like the loci within the vertebratemajor histocompatibility complex (MHC), Botryllus fusibility(or histocompatibility) genes are highly polymorphic. In Botryllus,the histocompatibility complex also controls self—sterility,and limits cross—fertilization between colonies sharinghistocompatibility alleles. The mouse MHC, the H-2 region, islinked to loci which also affect the frequencies of allelesat H-2 loci in mouse populations. Thus both systems containcharacters which could act to promote the heterozygous conditionat the linked histocompatibility loci. We suggest that suchlinked characters are responsible for the evolution of allogeneicpolymorphism in vertebrates (however currently maintained),and that tunicate fusibility loci may be the evolutionary precursorsof vertebrate MHC genes.     

Song Learning, Early Nutrition and Sexual Selection in Songbirds

SYNOPSIS. The developmental processes through which songbirdsacquire their species—typical songs have been well—studiedfrom a proximate perspective, but less attention has been givento the ultimate question of why birds learn to sing. We presenta new hypothesis for the adaptive significance of song learningin songbirds, suggesting that this specialized form of vocaldevelopment provides an indicator mechanism by which femalescan accurately assess the quality of potential mates. This hypothesisexpands on the established idea that song can provide an indicatorof male quality, but it explicitly links the variation in songexpression that females use to choose mates to the developmentalprocesses through which song is acquired. How well a male sings—reflectedin repertoire size or in other learned features of a male'ssinging behavior—provides an honest indicator of qualitybecause the timing of song learning and, more importantly, thetiming of the development of brain structures mediating learningcorresponds to a period in development during which young songbirdsare most likely to undergo nutritional stress. This correspondencemeans that song learning can provide a sensitive indicator ofearly developmental history in general, which in turn reflectsvarious aspects of the phenotypic and genotypic quality of apotential mate.     

The Ontogeny of Play within a Society: Preliminary Analysis

Data were collected on play behaviors occuring within a singletroop or rhesus monkeys in two environments. Observations wereregular and covered a total of almost 2 years. The three categoriesof play—object, activity, and social play—tendedto occur together, typically within the same recording session.Thus, the attending conditions—including satiation andmaturity—which permitted the occurrence of one categoryof play behavior, permitted the occurrence of the other categoriesas well. Maturity seemed to account for the appearance of aparticular behavior within a category (e.g., manipulation ratherthan touching; climbing rather than active hanging; rompingand wrestling rather than contact or touching). The social context,determined by such factors as maternal dominance, the identityof nearby animals, and the overall social tension of the troop,and reflected in the extent of inhibitory control of the targetanimals by their mothers, seemed to determine the frequencywith which these elements in the infant monkey's repertoirecould be displayed during any given period in the ontogeny ofthe individual.     

The evolutionary language game: an orthogonal approach

Evolutionary game dynamics have been proposed as a mathematical framework for the cultural evolution of language and more specifically the evolution of vocabulary. This article discusses a model that is mutually exclusive in its underlying principals with some previously suggested models. The model describes how individuals in a population culturally acquire a vocabulary by actively participating in the acquisition process instead of passively observing and communicate through peer-to-peer interactions instead of vertical parent-offspring relations. Concretely, a notion of social/cultural learning called the naming game is first abstracted using learning theory. This abstraction defines the required cultural transmission mechanism for an evolutionary process. Second, the derived transmission system is expressed in terms of the well-known selection-mutation model defined in the context of evolutionary dynamics. In this way, the analogy between social learning and evolution at the level of meaning-word associations is made explicit. Although only horizontal and oblique transmission structures will be considered, extensions to vertical structures over different genetic generations can easily be incorporated. We provide a number of simplified experiments to clarify our reasoning.     

Larval experience and latent effects--metamorphosis is not a new beginning

For many years ecologists have documented the remarkable within-speciesvariation inherent in natural systems—for example, variabilityin juvenile growth rates, mortality rates, fecundities, timeto reproductive maturity, the outcomes of competitive interactions,and tolerance to pollutants. Over the past 20 years, it hasbecome increasingly apparent that at least some of this variationmay reflect differences in embryonic or larval experiences.Such experiences may include delayed metamorphosis, short termstarvation, short term salinity stress, or exposure to sublethalconcentrations of pollutants or sublethal levels of ultra violetirradiation. Latent effects—effects that have their originsin early development but that are first exhibited in juvenilesor adults—have now been documented among gastropods, bivalves,echinoderms, polychaetes, crustaceans, bryozoans, urochordates,and vertebrates. The extent to which latent effects alter ecologicaloutcomes in natural populations in the field, and the mechanismsthrough which they are mediated are largely unexplored.     

Rapid adaptation: a new dimension for evolutionary perspectives in ecology

Although the study of adaptation is central to biology, two types of adaptation are recognized in the biological field: physiological adaptation (accommodation or acclimation; an individual organism’s phenotype is adjusted to its environment) and evolutionary–biological adaptation (adaptation is shaped by natural selection acting on genetic variation). The history of the former concept dates to the late nineteenth and early twentieth centuries, and has more recently been systemized in the twenty-first century. Approaches to the understanding of phenotypic plasticity and learning behavior have only recently been developed, based on cellular–histological and behavioral–neurobiological techniques as well as traditional molecular biology. New developments of the former concepts in phenotypic plasticity are discussed in bacterial persistence, wing di-/polymorphism with transgenerational effects, polyphenism in social insects, and defense traits for predator avoidance, including molecular biology analyses. We also discuss new studies on the concept of genetic accommodation resulting in evolution of phenotypic plasticity through a transgenerational change in the reaction norm based on a threshold model. Learning behavior can also be understood as physiological phenotypic plasticity, associating with the brain–nervous system, and it drives the accelerated evolutionary change in behavioral response (the Baldwin effect) with memory stock. Furthermore, choice behaviors are widely seen in decision-making of animal foragers. Incorporating flexible phenotypic plasticity and learning behavior into modeling can drastically change dynamical behavior of the system. Unification of biological sciences will be facilitated and integrated, such as behavioral ecology and behavioral neurobiology in the area of learning, and evolutionary ecology and molecular developmental biology in the theme of phenotypic plasticity.     

A Transactional Theory of Within-Group Conflict

Transactional models of social evolution emphasize that dominant members of the society can be favored to donate parcels of reproduction to subordinate members in return for cooperation. I construct a formal theory of intragroup conflict within the framework of transactional models by determining the maximum extent to which colony members can be selfish without destabilizing the group. The difference between the maximum value of the subordinate's fraction of group reproduction that the dominant can tolerate before ejecting the subordinate and the minimum value required by the subordinate to stay and cooperate peacefully in the group defines the "window of selfishness," which in turn predicts the frequency of within-group conflict. The window of selfishness tends to increase with increasing group reproductive output, increasingly harsh ecological constraints on solitary breeding, and, counterintuitively, increasing relatedness between subordinate and dominant. Increasing fighting ability of the subordinate can either widen or narrow the window of selfishness, the latter being most likely when ecological constraints on group living are strong. Although increasing relatedness is predicted to increase the rate of within-group aggression, the mean intensity of an aggressive act should decline, as predicted by the general theory of honest signaling between relatives and the tug-of-war models of within-group selfishness. In the bidding game, in which multiple dominants bid for the services of a subordinate, the window of selfishness is predicted to have zero width. A zero-width window of selfishness and low conflict also are predicted for saturated N-person groups, that is, groups whose total output is a concave function of group size and in which the dominant is not favored to admit additional subordinates. The model's predictions are compared to empirical evidence and to predictions of alternative models of intragroup aggression, including the value-aggression model and the pure tug-of-war model.     

TO AGE, TO DIE: PARITY, EVOLUTIONARY TRACKING AND COLE'S PARADOX

The existence of semelparity or "big bang" reproduction (reproducing only once in a lifetime) and iteroparity (reproducing more than once in a lifetime) has led to many questions investigating the evolution or persistence of these strategies. Here we investigate semelparity and iteroparity for their evolutionary importance. A mathematical model is used to illustrate how a population's ability to evolve depends on this life-history trait, and how this rate of evolution impacts the individual. We find that the ability of a trait to evolve is greater toward a semelparous strategy and this expresses a fitness advantage. This leads to an optimality between survival, population tracking ability, and lifetime fecundity.     

Mechanistic Approaches to Community Ecology: A New Reductionism

Mechanistic approaches to community ecology are those whichemploy individual— ecological concepts—those ofbehavioral ecology, physiological ecology, and ecomorphology—as theoretical bases for understanding community patterns. Suchapproaches, which began explicitly about a decade ago, are justnow coming into prominence. They stand in contrast to more traditionalapproaches, such as MacArthur and Levins (1967),which interpretcommunity ecology almost strictly in terms of "megaparameters.". Mechanistic approaches can be divided into those which use populationdynamics as a major component of the theory and those whichdo not; examples of the two are about equally common. The firstapproach sacrifices a highly detailed representation of individual—ecological processes; the second sacrifices an explicit representationof the abundance and persistence of populations. Three subdisciplines of ecology—individual, populationand community ecology—form a "perfect" hierarchy in Beckner's(1974) sense. Two other subdisciplines—ecosystem ecologyand evolutionary ecology—lie somewhat laterally to thishierarchy. The modelling of community phenomena using sets ofpopulation-dynamical equations is argued as an attempt at explanationvia the reduction of community to population ecology. Much ofthe debate involving Florida State ecologists is over whetheror not such a relationship is additive (or conjunctive), a verystrong form of reduction. I argue that reduction of communityto individual ecology is plausible via a reduction of populationecology to individual ecology. Approaches that derive the population-dynamicalequations used in population and community ecology from individual-ecologicalconsiderations, and which provide a decomposition of megaparametersinto behavioral and physiological parameters, are cited as illustratinghow the reduction might be done. I argue that "sufficient parameters"generally will not enhance theoretical understanding in communityecology. A major advantage of the mechanistic approach is that variationin population and community patterns can be understood as variationin individual-ecological conditions. In addition to enrichingthe theory, this allows the best functional form to be chosenfor modeling higher-level phenomena, where "best" is definedas biologically most appropriate rather than mathematicallymost convenient. Disadvantages of the mechanistic approach arethat it may portend an overly complex, massive and special theory,and that it naturally tends to avoid many-species phenomenasuch as indirect effects. The paper ends with a scenario fora mechanistic-ecological utopia.     

Ecological Bases of Hormone--Behavior Interactions: The "Emergency Life History Stage"

SYNOPSIS. Superimposed upon seasonal changes in morphology,physiology and behavior, are facultative responses to unpredictableevents known as labile (i.e., short-lived) perturbation factors(LPFs). These responses include behavioral and physiologicalchanges that enhance survival and collectively make up the "emergency"life history stage. There is considerable evidence that glucocorticosteroids,and other hormones in the hypothalamo—pituitary—adrenal(HPA) cascade, initiate and orchestrate the emergency life historystage within minutes to hours. This stage has a number of sub—stagesthat promote survival and avoid potential deleterious effectsof stress that may result from chronically elevated levels ofcirculating glucocorticosteroids over days and weeks. Thesesub—stages may include: redirection of behavior from anormal life history stage to increased foraging, irruptivetypemigration during the day, enhanced restfulness at night, andelevated gluconeogenesis. Once the perturbation passes, glucocorticosteroidsmay also promote recovery. Additional evidence from birds indicatesthat glucocorticosteroid responses to a standardized capture,handling and restraint protocol are modulated both on seasonaland individual levels. Field work reveals that these changesin responsiveness to LPFs have ecological bases, such as reproductivestate, body condition etc., that in turn indicate differenthormonal control mechanisms in the HPA cascade.