Inter-annual variability influences the eco-evolutionary dynamics of range-shifting

Peer reviewedPublisher PD

The speed of range shifts in fragmented landscapes

Peer reviewedPublisher PD

Rapid compensatory evolution promotes the survival of conjugative plasmids

Conjugative plasmids play a vital role in bacterial adaptation through horizontal gene transfer. Explaining how plasmids persist in host populations however is difficult, given the high costs often associated with plasmid carriage. Compensatory evolution to ameliorate this cost can rescue plasmids from extinction. In a recently published study we showed that compensatory evolution repeatedly targeted the same bacterial regulatory system, GacA/GacS, in populations of plasmid-carrying bacteria evolving across a range of selective environments. Mutations in these genes arose rapidly and completely eliminated the cost of plasmid carriage. Here we extend our analysis using an individual based model to explore the dynamics of compensatory evolution in this system. We show that mutations which ameliorate the cost of plasmid carriage can prevent both the loss of plasmids from the population and the fixation of accessory traits on the bacterial chromosome. We discuss how dependent the outcome of compensatory evolution is on the strength and availability of such mutations and the rate at which beneficial accessory traits integrate on the host chromosome

Bacteriophages limit the existence conditions for conjugative plasmids

Bacteriophages are a major cause of bacterial mortality and impose strong selection on natural bacterial populations, yet their effects on the dynamics of conjugative plasmids have rarely been tested. We combined experimental evolution, mathematical modeling, and individual-based simulations to explain how the ecological and population genetics effects of bacteriophages upon bacteria interact to determine the dynamics of conjugative plasmids and their persistence. The ecological effects of bacteriophages on bacteria are predicted to limit the existence conditions for conjugative plasmids, preventing persistence under weak selection for plasmid accessory traits. Experiments showed that phages drove faster extinction of plasmids in environments where the plasmid conferred no benefit, but they also revealed more complex effects of phages on plasmid dynamics under these conditions, specifically, the temporary maintenance of plasmids at fixation followed by rapid loss. We hypothesized that the population genetic effects of bacteriophages, specifically, selection for phage resistance mutations, may have caused this. Further mathematical modeling and individual-based simulations supported our hypothesis, showing that conjugative plasmids may hitchhike with phage resistance mutations in the bacterial chromosome

Spatial organisation of expanding bacterial colonies is affected by contact-dependent growth inhibition

Identifying how microbes are able to manipulate, survive and thrive in complex multispecies communities has expanded our understanding of how microbial ecosystems impact human health and the environment. The ability of bacteria to negatively affect neighbours, through explicit toxin delivery systems, provides them with an opportunity to manipulate the composition of growing microbial communities. Contact-dependent inhibition (CDI) systems (a Type Vb secretion system) are a distinct subset of competition systems whose contribution to shaping the development of spatially-structured bacterial communities are yet to be fully understood. Here we compare the impact of different CDI systems, at both the single cell and population level, to determine the key drivers of CDI-mediated competition within spatially-structured bacterial populations. Through an iterative approach using both an Escherichia coli experimental system and computational modelling, we show that CDI systems have subtle and system-specific effects at the single cell level, generating single cell wide boundaries between CDI-expressing inhibitor cells and their neighbouring targets. Despite the subtle effects of CDI at a single cell level, CDI systems greatly diminished the ability of susceptible targets to expand their range during colony growth. The inoculum density of the population, together with the CDI system-specific variables of the speed of inhibition after contact and biological cost of CDI, strongly affects CDI-mediated competition. In contrast, the magnitude of the toxin-induced growth retardation of target cells only weakly impacts the composition of the population. Our work reveals how distinct CDI systems can differentially affect the composition and spatial arrangement of bacterial populations

Landscape as a Model: The Importance of Geometry

In all models, but especially in those used to predict uncertain processes (e.g., climate change and nonnative species establishment), it is important to identify and remove any sources of bias that may confound results. This is critical in models designed to help support decisionmaking. The geometry used to represent virtual landscapes in spatially explicit models is a potential source of bias. The majority of spatial models use regular square geometry, although regular hexagonal landscapes have also been used. However, there are other ways in which space can be represented in spatially explicit models. For the first time, we explicitly compare the range of alternative geometries available to the modeller, and present a mechanism by which uncertainty in the representation of landscapes can be incorporated. We test how geometry can affect cell-to-cell movement across homogeneous virtual landscapes and compare regular geometries with a suite of irregular mosaics. We show that regular geometries have the potential to systematically bias the direction and distance of movement, whereas even individual instances of landscapes with irregular geometry do not. We also examine how geometry can affect the gross representation of real-world landscapes, and again show that individual instances of regular geometries will always create qualitative and quantitative errors. These can be reduced by the use of multiple randomized instances, though this still creates scale-dependent biases. In contrast, virtual landscapes formed using irregular geometries can represent complex real-world landscapes without error. We found that the potential for bias caused by regular geometries can be effectively eliminated by subdividing virtual landscapes using irregular geometry. The use of irregular geometry appears to offer spatial modellers other potential advantages, which are as yet underdeveloped. We recommend their use in all spatially explicit models, but especially for predictive models that are used in decisionmaking

Differing effects of age and starvation on reproductive performance in Drosophila melanogaster

Successful reproduction requires the completion of many, often condition-dependent, stages, from mate searching and courtship through to sperm transfer, fertilisation and offspring production. Animals can plastically adjust their investment in each stage according to the physical and social environment, their own condition, their future reproductive potential, and the condition of their partner. Here we manipulate age and condition, through a nutritional challenge early or late in life, of both male and female Drosophila melanogaster and measure the effects on courtship, mating, and fitness when paired with a standardized (unmanipulated) partner. Older males were slower to start courting and mating, and courted at a slower rate, but males were indifferent to female age or condition despite older females laying and hatching fewer eggs. Female condition had a substantial effect on mating acceptance rate, which dropped dramatically after starvation, and particularly recent starvation experience. In contrast, male condition had little effect on any of the components of reproductive performance we measured. Intriguingly, we found no evidence for additive or multiplicative effects of ageing and starvation: the only significant interaction between these variables was on male latency to initiate courtship - older males were slower to start courting unless they had experienced starvation early in life. These results indicate that the immediate costs of mating differ between males and females, and that the sexes differ in their perception of the opportunity cost sustained by refusing a mating opportunity. Our results support the idea that ageing has more wide-ranging impact on reproductive behaviours than does nutritional challenge

Climate change, climatic variation and extreme biological responses

Extreme climatic events could be major drivers of biodiversity change, but it is unclear whether extreme biological changes are (i) individualistic (species- or group-specific), (ii) commonly associated with unusual climatic events and/or (iii) important determinants of long-term population trends. Using population time series for 238 widespread species (207 Lepidoptera and 31 birds) in England since 1968, we found that population 'crashes' (outliers in terms of species' year-to-year population changes) were 46% more frequent than population 'explosions'. (i) Every year, at least three species experienced extreme changes in population size, and in 41 of the 44 years considered, some species experienced population crashes while others simultaneously experienced population explosions. This suggests that, even within the same broad taxonomic groups, species are exhibiting individualistic dynamics, most probably driven by their responses to different, short-term events associated with climatic variability. (ii) Six out of 44 years showed a significant excess of species experiencing extreme population changes (5 years for Lepidoptera, 1 for birds). These 'consensus years' were associated with climatically extreme years, consistent with a link between extreme population responses and climatic variability, although not all climatically extreme years generated excess numbers of extreme population responses. (iii) Links between extreme population changes and long-term population trends were absent in Lepidoptera and modest (but significant) in birds. We conclude that extreme biological responses are individualistic, in the sense that the extreme population changes of most species are taking place in different years, and that long-term trends of widespread species have not, to date, been dominated by these extreme changes.This article is part of the themed issue 'Behavioural, ecological and evolutionary responses to extreme climatic events'

Theoretical and methodological approaches to ecological changes, social behaviour and human intergroup tolerance 300,000 to 30,0000bp

Archaeological evidence suggests that important shifts were taking place in the character of human social behaviours 300,000 to 30,000 years ago. New artefact types appear and are disseminated with greater frequency. Transfers of both raw materials and finished artefacts take place over increasing distances, implying larger scales of regional mobility and more frequent and friendlier interactions between different communities. While these changes occur during a period of increasing environmental variability, the relationship between ecological changes and transformations in social behaviours is elusive. Here, we explore a possible theoretical approach and methodology for understanding how ecological contexts can influence selection pressures acting on intergroup social behaviours. We focus on the relative advantages and disadvantages of intergroup tolerance in different ecological contexts using agent-based modelling (ABM). We assess the relative costs and benefits of different 'tolerance' levels in between-group interactions on survival and resource exploitation in different environments. The results enable us to infer a potential relationship between ecological changes and proposed changes in between-group behavioural dynamics. We conclude that increasingly harsh environments may have driven changes in hormonal and emotional responses in humans leading to increasing intergroup tolerance, i.e. transformations in social behaviour associated with ‘self-domestication’. We argue that changes in intergroup tolerance is a more parsimonious explanation for the emergence of what has been seen as ‘modern human behaviour’ than changes in hard aspects of cognition or other factors such as cognitive adaptability or population size

Living to fight another day : The ecological and evolutionary significance of Neanderthal healthcare

Evidence of care for the ill and injured amongst Neanderthals, inferred through skeletal evidence for survival from severe illness and injury, is widely accepted. However, healthcare practices have been viewed primarily as an example of complex cultural behaviour, often discussed alongside symbolism or mortuary practices. Here we argue that care for the ill and injured is likely to have a long evolutionary history and to have been highly effective in improving health and reducing mortality risks. Healthcare provisioning can thus be understood alongside other collaborative ‘risk pooling’ strategies such as collaborative hunting, food sharing and collaborative parenting. For Neanderthals in particular the selective advantages of healthcare provisioning would have been elevated by a variety of ecological conditions which increased the risk of injury as well their particular behavioural adaptations which affected the benefits of promoting survival from injury and illness. We argue that healthcare provisioning was not only a more significant evolutionary adaptation than has previously been acknowledged, but moreover may also have been essential to Neanderthal occupation at the limits of the North Temperate Zone