Evolution in a Changing Environment

We propose a simple model for genetic adaptation to a changing environment, describing a fitness landscape characterized by two maxima. One is associated with “specialist” individuals that are adapted to the environment; this maximum moves over time as the environment changes. The other maximum is static, and represents “generalist” individuals not affected by environmental changes. The rest of the landscape is occupied by “maladapted” individuals. Our analysis considers the evolution of these three subpopulations. Our main result is that, in presence of a sufficiently stable environmental feature, as in the case of an unchanging aspect of a physical habitat, specialists can dominate the population. By contrast, rapidly changing environmental features, such as language or cultural habits, are a moving target for the genes; here, generalists dominate, because the best evolutionary strategy is to adopt neutral alleles not specialized for any specific environment. The model we propose is based on simple assumptions about evolutionary dynamics and describes all possible scenarios in a non-trivial phase diagram. The approach provides a general framework to address such fundamental issues as the Baldwin effect, the biological basis for language, or the ecological consequences of a rapid climate change.     

Helical Antifreeze Proteins Have Independently Evolved in Fishes on Four Occasions

Alanine-rich α-helical (type I) antifreeze proteins (AFPs) are produced by a variety of fish species from three different orders to protect against freezing in icy seawater. Interspersed amongst and within these orders are fishes making AFPs that are completely different in both sequence and structure. The origin of this variety of types I, II, III and antifreeze glycoproteins (AFGPs) has been attributed to adaptation following sea-level glaciations that occurred after the divergence of most of the extant families of fish. The presence of similar types of AFPs in distantly related fishes has been ascribed to lateral gene transfer in the case of the structurally complex globular type II lectin-like AFPs and to convergent evolution for the AFGPs, which consist of a well-conserved tripeptide repeat. In this paper, we examine the genesis of the type I AFPs, which are intermediate in complexity. These predominantly α-helical peptides share many features, such as putative capping structures, Ala-richness and amphipathic character. We have added to the type I repertoire by cloning additional sequences from sculpin and have found that the similarities between the type I AFPs of the four distinct groups of fishes are not borne out at the nucleotide level. Both the non-coding sequences and the codon usage patterns are strikingly different. We propose that these AFPs arose via convergence from different progenitor helices with a weak affinity for ice and that their similarity is dictated by the propensity of specific amino acids to form helices and to align water on one side of the helix into an ice-like pattern.     

Evolved psychology in a novel environment

The human “environment of evolutionary adaptedness” can only be inferred indirectly. In contrast, the behavior of some nonhuman animals can be compared among “natural” and various altered environments. As an example, male immigration tactics in unprovisioned versus provisioned macaque (Macaca) populations are compared using Tooby and Cosmides’s (1992) framework for evolutionary functional analysis. In unprovisioned populations, social groups contain few males, and immigrant male takeovers of alpha rank occur frequently. In provisioned populations, groups contain many males, and males almost invariably enter social groups at very low rank and rise in rank only as more dominant males emigrate or die. Male conformity to the “seniority rule” is hypothesized to represent the behavioral output of an evolved decision-making algorithm (psychological mechanism) that takes into account (1) the net payoff of each rank in the dominance hierarchy and (2) the power of male group size as a predictor of the likelihood of successful immigrant takeover. Joseph H. Manson is Assistant Professor of Anthropology at the University of California, Los Angeles. His research interests are social relationships in nonhuman primates and humans, with particular emphases on mate choice, courtship tactics, intrasexual competition, and (currently) mother-infant relationships and infant handling. He has conducted fieldwork on rhesus macaques at Cayo Santiago and white-faced capuchins in Costa Rica.     

Nonplastid Eukaryotic Response Regulators Have a Monophyletic Origin and Evolved from Their Bacterial Precursors in Parallel with Their Cognate Sensor Kinases

We have analyzed all currently sequenced eukaryotic proteins containing either a kinase module or a receiver module, corresponding to those found in bacterial sensor kinases or response regulators, respectively, of the so-called two-component regulatory systems. We demonstrate that the eukaryotic receiver modules belong to a single subfamily of the bacterial receiver modules. Moreover, the cognate eukaryotic kinase modules exhibit a similar clustering pattern on the sensor kinase phylogenetic tree, suggesting that they evolved in parallel with the receiver modules from a common ancestral source that bore both modules. Multiple alignments of the sequences corresponding to these modules are presented and discussed, and eukaryotic-specific signature sequences are derived. Received: 18 October 1995 / Accepted: 16 December 1996     

Could Have Gone Wrong: Effects of Abrupt Changes in Migratory Behaviour on Harvest in a Waterbird Population

To sustainably exploit a population, it is crucial to understand and reduce uncertainties about population processes and effects of harvest. In migratory species, management is challenged by geographically separated changing environmental conditions, which may cause unexpected changes in species distribution and harvest. We describe the development in the harvest of Svalbard-breeding pink-footed geese (Anser brachyrhynchus) in relation to the observed trajectory and migratory behaviour of the population. In autumn, geese migrate via stopover sites in Norway and Denmark (where they are hunted) to wintering grounds in the Netherlands and Belgium (where they are protected). In Denmark and Norway harvesting increased stepwise during the 2000s. The increase in the population size only partly explained the change. The change corresponded to a simultaneous stepwise increase in numbers of geese staging in Denmark throughout autumn and winter; geese also moved further inland to feed which collectively increased their exposure to hunting. In Norway the increase in harvest reflected greater utilisation of lowland farmland areas by geese, increasing their hunting exposure. The study demonstrates how changes in migratory behaviour can abruptly affect exposure to hunting, which showed a functional response to increased temporal and spatial availability of geese. The harvest has now reached a level likely to cause a population decline. It highlights the need for flexible, internationally coordinated hunting regulations and reliable up-to-date population estimates and hunting bag statistics, which are rare in European management of migratory waterbirds. Without such information decisions are left with judgments based on population estimates, which often have time lags of several years between recording and reporting, hampering possibilities for the timely adjustment of management actions.     

Species Management Benchmarking: Outcomes Over Outputs in a Changing Operating Environment

Species management has been utilized by the zoo and aquarium industry, since the mid‐1990s, to ensure the ongoing genetic and demographic viability of populations, which can be difficult to maintain in the ever‐changing operating environments of zoos. In 2009, the Zoo and Aquarium Association Australasia reviewed their species management services, focusing on addressing issues that had arisen as a result of the managed programs maturing and operating environments evolving. In summary, the project examined resourcing, policies, processes, and species to be managed. As a result, a benchmarking tool was developed (Health Check Report, HCR), which evaluated the programs against a set of broad criteria. A comparison of managed programs (n = 98), between 2008 and 2011, was undertaken to ascertain the tool's effectiveness. There was a marked decrease in programs that were designated as weak (37 down to 13); and an increase in excellent programs (24 up to 49) between the 2 years. Further, there were significant improvements in the administration benchmarking area (submission of reports, captive management plan development) across a number of taxon advisory groups. This HCR comparison showed that a benchmarking tool enables a program's performance to be quickly assessed and any remedial measures applied. The increases observed in program health were mainly due to increased management goals being attained. The HCR will be an ongoing program, as the management of the programs increases and goals are achieved, criteria will be refined to better highlight ongoing issues and ways in which these can be resolved. Zoo Biol. 32:230–237, 2013. © 2012 Wiley Periodicals, Inc.     

Leaflet Movement of Robinia pseudoacacia in Response to a Changing Light Environment

Diurnal and nocturnal leaflet movement of black locust （Robinia pseudoacacia L.） was investigated under three light schemes： 100% natural irradiance, 50% shading, and 90% shading. Changes in leaf mid-vein angle were described by measurements of two planes： （i） β, the angle formed by the bottom of the petiolule and its relation to the horizontal plane; and （ii） θ, the angle between the petiolule and the main leaflet vein. The two highest light regimens had a significant effect on β. Variation in β tends to make the leaflet more erect, thereby minimizing any negative impact of high irradiance on leaf lamina. Light-dark rhythms induced variation in θ （termed nyctinastic movement）. Nyctinastic movement is important during the low light levels experienced by leaflets in early morning and late afternoon. At low light levels, the leaflet stopped nyctinastic movement and θ was fixed at an angle that may have enabled the leaf lamina to maximize light interception. After the light-dark cycle was reestablished, nyctinastic movement was restored. Taken together, our results suggest that irradiance induces variation in β leading to diurnal leaflet movement （diaheliotropism）, whereas the light-dark cycle influences θ, which results in nocturnal leaflet movement. Both angles are important for describing patterns of leaf movement in R. pseudocacia.     

Modelling Plant Growth Under Varying Environment Conditions in a Uniform Canopy

A model for the growth of plants is proposed. It extends theconcept of thermal time to include the effects of photosyntheticallyactive radiation and carbon dioxide concentration in a formulathat is analogous to the rate of flow through a set of conductancesin series. The formula transforms time periods of differingor varying environmental conditions to a consistent plant-relatedbasis. The integrated plant environmental time is used in theexpolinear equation of Gourdriaan and Monteith (1990) to providea simple dynamic model for the growth of mutually shading plantsin a uniform canopy of an even-aged monoculture. The model is fitted to data from lettuce and cauliflower. Eachhad been grown, in a glasshouse, at four different times ofyear and at four densities. The plant environmental time relationallows for the different conditions of temperatures and light,and the expolinear equation then predicts the set of growthcurves at the different densities. Parameters values differbetween the species, as would the predicted values of environmentaltime. An improved fit is obtained by using the mutual shading relationto modify the radiation term only, rather than the whole environmentalgrowth rate expression. Most of the observed results differby less than one day in time from the fitted values. Corroborationfor the model is provided by values of an effective leaf arearatio, estimated from the effects of plant density and plantweight upon competition for radiation.Copyright 1993, 1999 AcademicPress Growth, model, temperature, PAR, carbon dioxide, plant environmental time, density, expolinear, monocrop, competition, lettuce, cauliflower     

Early Olfactory Environment Influences Social Behaviour in Adult Octodon degus

We evaluated the extent to which manipulation of early olfactory environment can influence social behaviours in the South American Hystricognath rodent Octodon degus. The early olfactory environment of newborn degus was manipulated by scenting all litter members with eucalyptol during the first month of life. The social behaviour of sexually mature animals (5–7 months old) towards conspecifics was then assessed using a y-maze to compare the response of control (naïve) and treated animals to two different olfactory configurations (experiment 1): (i) a non-familiarized conspecific impregnated with eucalyptol (eucalyptol arm) presented against (ii) a non-familiarized unscented conspecific (control arm). In addition, in dyadic encounters, we assessed the behaviour of control and eucalyptol treated animals towards a non-familiarized conspecific scented with eucalyptol (experiment 2). We found that control subjects explored and spent significantly less time in the eucalyptol arm, indicating neophobic behaviours towards the artificially scented conspecific. Treated subjects explored and spent similar time in both arms of the maze, showing the same interest for both olfactory stimuli presented. During dyadic encounters in experiment 2, an interaction effect between early experience and sex was observed. Control males escaped and avoided their scented partner more frequently than eucalyptol treated male subjects and than females. Both groups did not differ in the exploration of their scented partners, suggesting that avoidance within agonistic context does not relate to neophobic behaviours. Our results suggest that the exposure to eucalyptol during early ontogeny decreases evasive behaviours within an agonistic context as a result of olfactory learning. Altogether, these results indicate that olfactory cues learned in early ontogeny can influence olfactory-guided behaviours in adult degus.     

Cassava Production and Pest Management: Present and Potential Threats in a Changing Environment

Cassava is attacked by a complex of arthropod pests across the tropical regions of the world where the crop is grown. Root yield losses have been recorded for several pests, including mites, mealybugs, whiteflies, hornworm, lacebugs, thrips and burrower bugs. Agronomic characteristics such as vegetative propagation, a long growth cycle, drought tolerance, staggered planting dates and intercropping contribute to the considerable diversity of pests that feed on the crop. The dynamics of cassava production are evolving as trends in the food, feed and industrial starch sector are leading to an increased demand for high quality starches. The resulting shift to larger scale production units, expansion of cultivated area and modifications in crop management combined with the effects of climate change, especially warmer temperatures and altered rainfall patterns, affect the occurrence and dynamics of arthropod pests in cassava agro ecosystems. Data is presented to describe the effects of temperature and dry seasons on key pest species. Whiteflies, mites and mealybugs register a suitability increase in the same areas in South America: Northeastern Brazil, Northern Argentina, South-Central Bolivia, and Southwest Peru. In Africa increases are projected in Southeast Africa and Madagascar. In Asia, regions with greater projected suitability for these pest species are Coastal India and Southeast Asia. Future trends and important criteria that will influence the severity and management of key pests are discussed.     

Conditions Under Which Nitrogen Can Limit Steady-State Net Primary Production in a General Class of Ecosystem Models

Human activity is drastically altering global nitrogen (N) availability. The extent to which ecosystems absorb additional N—and with it, additional CO₂—depends on whether net primary production (NPP) is N-limited, so it is important to understand conditions under which N can limit NPP. Here I use a general dynamical model to show that N limitation at steady-state—such as in old-growth forests—depends on the balance of biotically controllable versus uncontrollable N inputs and losses. Steady-state N limitation is only possible when uncontrollable inputs (for example, atmospheric deposition) exceed controllable losses (for example, leaching of plant-available soil N), which is the same as when uncontrollable losses (for example, leaching of plant-unavailable soil N) exceed controllable inputs (biological N fixation). These basic results are robust to many model details, such as the number of plant-unavailable soil N pools and the number and type of N fixers. Empirical data from old-growth tropical (Hawai’i) and temperate (Oregon, Washington, Chile) forests support the model insights. Practically, this means that any N fixer—symbiotic or not—could overcome ecosystem N limitation, so understanding N limitation requires understanding controls on all N fixers. Further, comparing losses of plant-available N to abiotic inputs could offer a rapid diagnosis of whether ecosystems can be N-limited, although the applicability of this result is constrained to ecosystems with a steady-state N cycle such as old-growth forests largely devoid of disturbance.     

Frogs Have Rules: Selective Attention Algorithms Regulate Chorusing in Physalaemus pustulosus (Leptodactylidae)

In various acoustic insects and frogs, females preferentially orient towards the leading of two or more males' advertisement signals that occur closely in time. Such preferences in receivers have apparently selected for timing mechanisms whereby male signallers actively refrain from calling immediately following the onset of a neighbour's call and thereby increase their production of leading calls. However, indiscriminate application of this inhibitory mechanism to all neighbours might severely reduce a male's calling rate, particularly in high density. Consequently, mechanisms of selective attention to only a subset of signalling neighbours are expected.
Female Túngara frogs ( Physalaemus pustulosus ) exhibit strong preferences for leading male calls, and males refrain from calling immediately following a neighbour. Four-loudspeaker playback experiments demonstrated that males do selectively apply this inhibitory mechanism to only a subset of close signalling neighbours. Selective attention is regulated by a combination of sliding threshold and fixed number rules: (i) Attend to the loudest (nearest) conspecific neighbour and those additional ones whose calls are within 6–8 dB of the loudest one; (ii) attend to only two neighbours in total when the calls are weak or the second one is much farther than the first; (iii) attend to three neighbours when the calls are loud or all neighbours are approximately equidistant. The means by which such plasticity may be achieved and its potential adaptiveness are discussed.     

Ensembles of Spiking Neurons with Noise Support Optimal Probabilistic Inference in a Dynamically Changing Environment

It has recently been shown that networks of spiking neurons with noise can emulate simple forms of probabilistic inference through “neural sampling”, i.e., by treating spikes as samples from a probability distribution of network states that is encoded in the network. Deficiencies of the existing model are its reliance on single neurons for sampling from each random variable, and the resulting limitation in representing quickly varying probabilistic information. We show that both deficiencies can be overcome by moving to a biologically more realistic encoding of each salient random variable through the stochastic firing activity of an ensemble of neurons. The resulting model demonstrates that networks of spiking neurons with noise can easily track and carry out basic computational operations on rapidly varying probability distributions, such as the odds of getting rewarded for a specific behavior. We demonstrate the viability of this new approach towards neural coding and computation, which makes use of the inherent parallelism of generic neural circuits, by showing that this model can explain experimentally observed firing activity of cortical neurons for a variety of tasks that require rapid temporal integration of sensory information.     

Rapid Establishment of a Regular Distribution of Adult Tropical Drosophila Parasitoids in a Multi-Patch Environment by Patch Defence Behaviour

Females of the larval parasitoid of Drosophila, Asobara citri, from sub-Saharan Africa, defend patches with hosts by fighting and chasing conspecific females upon encounter. Females of the closely related, palearctic species Asobara tabida do not defend patches and often search simultaneously in the same patch. The effect of patch defence by A. citri females on their distribution in a multi-patch environment was investigated, and their distributions were compared with those of A. tabida. For both species 20 females were released from two release-points in replicate experiments. Females of A. citri quickly reached a regular distribution across 16 patches, with a small variance/mean ratio per patch. Conversely, A. tabida females initially showed a clumped distribution, and after gradual dispersion, a more Poisson-like distribution across patches resulted (variance/mean ratio was closer to 1 and higher than for A. citri). The dispersion of A. tabida was most probably an effect of exploitation: these parasitoids increasingly made shorter visits to already exploited patches. We briefly discuss hypotheses on the adaptive significance of patch defence behaviour or its absence in the light of differences in the natural history of both parasitoid species, notably the spatial distribution of their hosts.     

Social Behaviour and Cooperative Breeding in Arctic Foxes,Alopex lagopus (L.), in a Semi-natural Environment

Most canid species show cooperative breeding at least occasionally. The helper-at-the-den system, when extra adults serve as helpers by feeding and guarding the cubs of an alpha pair, has been observed but not studied in any detail in wild Arctic foxes (Alopex lagopus). During a 3-months study of arctic foxes in two enclosures of 4 ha each, we measured the social behaviour during the reproductive season. Older foxes dominated younger ones and males dominated females of the same age. A litter with one surviving cub was born in one enclosure. The alpha male increased his rate of urine marking and barking and fed the alpha female both before and after the birth of the litter. However, about 10 days after the birth, the alpha female died. The cub was fed by his putative father, his sister and his brother (both one year old). The one year old female increased her rate of territorial defence, measured as urine marking and barking, when the mother died. The subordinate females were probably suppressed from breeding by the high aggression levels and territorial defence of the dominant females in each enclosure. The dominant female in the second enclosure came into heat after the death of the alpha female (her mother) in the first enclosure. These changes in behaviour can probably be explained by sexual inhibition by the alpha female while she was present. The significance of territorial defence and dominance, inbreeding avoidance, sexual suppression and evolution of helping behaviour are discussed.     

A Few Bad Apples: A Model of Disease Influenced Agent Behaviour in a Heterogeneous Contact Environment

For diseases that infect humans or livestock, transmission dynamics are at least partially dependent on human activity and therefore human behaviour. However, the impact of human behaviour on disease transmission is relatively understudied, especially in the context of heterogeneous contact structures such as described by a social network. Here, we use a strategic game, coupled with a simple disease model, to investigate how strategic agent choices impact the spread of disease over a contact network. Using beliefs that are based on disease status and that build up over time, agents choose actions that stochastically determine disease spread on the network. An agent’s disease status is therefore a function of both his own and his neighbours actions. The effect of disease on agents is modelled by a heterogeneous payoff structure. We find that the combination of network shape and distribution of payoffs has a non-trivial impact on disease prevalence, even if the mean payoff remains the same. An important scenario occurs when a small percentage (called noncooperators) have little incentive to avoid disease. For diseases that are easily acquired when taking a risk, then even when good behavior can lead to disease eradication, a small increase in the percentage of noncooperators (less than 5%) can yield a large (up to 25%) increase in prevalence.     

Trait Values,Not Trait Plasticity,Best Explain Invasive Species' Performance in a Changing Environment

The question of why some introduced species become invasive and others do not is the central puzzle of invasion biology. Two of the principal explanations for this phenomenon concern functional traits: invasive species may have higher values of competitively advantageous traits than non-invasive species, or they may have greater phenotypic plasticity in traits that permits them to survive the colonization period and spread to a broad range of environments. Although there is a large body of evidence for superiority in particular traits among invasive plants, when compared to phylogenetically related non-invasive plants, it is less clear if invasive plants are more phenotypically plastic, and whether this plasticity confers a fitness advantage. In this study, I used a model group of 10 closely related Pinus species whose invader or non-invader status has been reliably characterized to test the relative contribution of high trait values and high trait plasticity to relative growth rate, a performance measure standing in as a proxy for fitness. When grown at higher nitrogen supply, invaders had a plastic RGR response, increasing their RGR to a much greater extent than non-invaders. However, invasive species did not exhibit significantly more phenotypic plasticity than non-invasive species for any of 17 functional traits, and trait plasticity indices were generally weakly correlated with RGR. Conversely, invasive species had higher values than non-invaders for 13 of the 17 traits, including higher leaf area ratio, photosynthetic capacity, photosynthetic nutrient-use efficiency, and nutrient uptake rates, and these traits were also strongly correlated with performance. I conclude that, in responding to higher N supply, superior trait values coupled with a moderate degree of trait variation explain invasive species'' superior performance better than plasticity per se.