Integrodifference models for evolutionary processes in biological invasions.

Individual variability in dispersal and reproduction abilities can lead to evolutionary processes that may have significant effects on the speed and shape of biological invasions. Spatial sorting, an evolutionary process through which individuals with the highest dispersal ability tend to agglomerate at the leading edge of an invasion front, and spatial selection, spatially heterogeneous forces of selection, are among the fundamental evolutionary forces that can change range expansions. Most mathematical models for these processes are based on reaction-diffusion equations, i.e., time is continuous and dispersal is Gaussian. We develop novel theory for how evolution shapes biological invasions with integrodifference equations, i.e., time is discrete and dispersal can follow a variety of kernels. Our model tracks how the distribution of growth rates and dispersal ability in the population changes from one generation to the next in continuous space. We include mutation between types and a potential trade-off between dispersal ability and growth rate. We perform the analysis of such models in continuous and discrete trait spaces, i.e., we determine the existence of travelling wave solutions, asymptotic spreading speeds and their linear determinacy, as well as the population distributions at the leading edge. We also establish the relation between asymptotic spreading speeds and mutation probabilities. We observe conditions for when spatial sorting emerges and when it does not and also explore conditions where anomalous spreading speeds occur, as well as possible effects of deleterious mutations in the population.

Cortical Electrocorticogram (ECoG) Is a Local Signal.

Electrocorticogram (ECoG), obtained by low-pass filtering the brain signal recorded from a macroelectrode placed on the cortex, is extensively used to find the seizure focus in drug-resistant epilepsy and is of growing importance in cognitive and brain-machine-interfacing studies. To accurately estimate the epileptogenic cortex or to make inferences about cognitive processes, it is important to determine the "spatial spread" of ECoG (i.e., the extent of cortical tissue that contributes to its activity). However, the ECoG spread is currently unknown; even the spread of local field potential (LFP) obtained from microelectrodes is debated, with estimates ranging from a few hundred micrometers to several millimeters. Spatial spread can be estimated by measuring the receptive field (RF) and multiplying by the cortical magnification factor, but this method overestimates the spread because RF size gets inflated due to several factors. This issue can be partially addressed using a model that compares the RFs of two measures, such as LFP and multi-unit activity (MUA). To use this approach for ECoG, we designed a customized array containing both microelectrodes and ECoG electrodes to simultaneously map MUA, LFP, and ECoG RFs from the primary visual cortex of awake monkeys (three female Macaca radiata). The spatial spread of ECoG was surprisingly local (diameter ∼3 mm), only 3 times that of the LFP. Similar results were obtained using a model to simulate ECoG as a sum of LFPs of varying electrode sizes. Our results further validate the use of ECoG in clinical and basic cognitive research.SIGNIFICANCE STATEMENT Brains signals capture different attributes of the neural network depending on the size and location of the recording electrode. Electrocorticogram (ECoG), obtained by placing macroelectrodes (typically 2-3 mm diameter) on the exposed surface of the cortex, is widely used by neurosurgeons to identify the source of seizures in drug-resistant epileptic patients. The brain area responsible for seizures is subsequently surgically removed. Accurate estimation of the epileptogenic cortex and its removal requires the estimation of spatial spread of ECoG. Here, we estimated the spatial spread of ECoG in five behaving monkeys using two different approaches. Our results suggest that ECoG is a local signal (diameter of ∼3 mm), which can provide a useful tool for clinical, cognitive neuroscience, and brain-machine-interfacing applications.

Community context and dispersal stochasticity drive variation in spatial spread.

Dispersal is a key process in shaping species spatial distributions. Species interactions and variation in dispersal probabilities may jointly influence species spatial dynamics. However, many studies examine dispersal as a neutral process, independent of community context or intraspecific variation in dispersal behaviour. Here, we use controlled, replicated communities of two Tribolium species (T. castaneum and T. confusum) to examine how intraspecific variation in dispersal behaviour and community context influence dispersal dynamics in simple experimental landscapes composed of homogeneous habitat patches. We found considerable individual-level variation in dispersal probability that was unrelated to body size variation. Further, the context of dispersal mattered, as T. castaneum dispersal was reduced in two-species communities, while T. confusum dispersal was unaffected by community composition. Incorporating individual-level variation into a two-species stochastic spatial Ricker model, we provide evidence that individual-level variability in dispersal behaviour results in more variable spatial spread than assuming individuals have the same dispersal probability. Further, interspecific competition resulted in more variable spatial spread. The variability in spatial spread observed in our tightly controlled and replicated experimental system and in our stochastic model simulations points to potential fundamental limitations in forecasting species shifting ranges without considering potential interspecific interactions and demographic variability in dispersal behaviour.

Local translation in neuronal compartments: how local is local?

Efficient neuronal function depends on the continued modulation of the local neuronal proteome. Local protein synthesis plays a central role in tuning the neuronal proteome at specific neuronal regions. Various aspects of translation such as the localization of translational machinery, spatial spread of the newly translated proteins, and their site of action are carried out in specialized neuronal subcompartments to result in a localized functional outcome. In this review, we focus on the various aspects of these local translation compartments such as size, biochemical and organelle composition, structural boundaries, and temporal dynamics. We also discuss the apparent absence of definitive components of translation in these local compartments and the emerging state-of-the-art tools that could help dissecting these conundrums in greater detail in the future.

How social structures, space, and behaviors shape the spread of infectious diseases using chikungunya as a case study.

Whether an individual becomes infected in an infectious disease outbreak depends on many interconnected risk factors, which may relate to characteristics of the individual (e.g., age, sex), his or her close relatives (e.g., household members), or the wider community. Studies monitoring individuals in households or schools have helped elucidate the determinants of transmission in small social structures due to advances in statistical modeling; but such an approach has so far largely failed to consider individuals in the wider context they live in. Here, we used an outbreak of chikungunya in a rural community in Bangladesh as a case study to obtain a more comprehensive characterization of risk factors in disease spread. We developed Bayesian data augmentation approaches to account for uncertainty in the source of infection, recall uncertainty, and unobserved infection dates. We found that the probability of chikungunya transmission was 12% [95% credible interval (CI): 8-17%] between household members but dropped to 0.3% for those living 50 m away (95% CI: 0.2-0.5%). Overall, the mean transmission distance was 95 m (95% CI: 77-113 m). Females were 1.5 times more likely to become infected than males (95% CI: 1.2-1.8), which was virtually identical to the relative risk of being at home estimated from an independent human movement study in the country. Reported daily use of antimosquito coils had no detectable impact on transmission. This study shows how the complex interplay between the characteristics of an individual and his or her close and wider environment contributes to the shaping of infectious disease epidemics.

Sex difference and Allee effects shape the dynamics of sex-structured invasions.

The rate at which a population grows and spreads can depend on individual behaviour and interactions with others. In many species with two sexes, males and females differ in key life-history traits (e.g. growth, survival and dispersal), which can scale up to affect population rates of growth and spread. In sexually reproducing species, the mechanics of locating mates and reproducing successfully introduce further complications for predicting the invasion speed (spread rate), as both can change nonlinearly with density. Most models of population spread are based on one sex, or include limited aspects of sex differences. Here we ask whether and how the dynamics of finding mates interact with sex-specific life-history traits to influence the rate of population spread. We present a hybrid approach for modelling invasions of populations with two sexes that links individual-level mating behaviour (in an individual-based model) to population-level dynamics (in an integrodifference equation model). We find that limiting the amount of time during which individuals can search for mates causes a demographic Allee effect which can slow, delay, or even prevent an invasion. Furthermore, any sex-based asymmetries in life history or behaviour (skewed sex ratio, sex-biased dispersal, and sex-specific mating behaviours) amplify these effects. In contrast, allowing individuals to mate more than once ameliorates these effects, enabling polygynandrous populations to invade under conditions where monogamously mating populations would fail to establish. We show that details of individuals' mating behaviour can impact the rate of population spread. Based on our results, we propose a stricter definition of a mate-finding Allee effect, which is not met by the commonly used minimum mating function. Our modelling approach, which links individual- and population-level dynamics in a single model, may be useful for exploring other aspects of individual behaviour that are thought to impact the rate of population spread.

Travel distance and human movement predict paths of emergence and spatial spread of chikungunya in Thailand.

Human movement contributes to the probability that pathogens will be introduced to new geographic locations. Here we investigate the impact of human movement on the spatial spread of Chikungunya virus (CHIKV) in Southern Thailand during a recent re-emergence. We hypothesised that human movement, population density, the presence of habitat conducive to vectors, rainfall and temperature affect the transmission of CHIKV and the spatiotemporal pattern of cases seen during the emergence. We fit metapopulation transmission models to CHIKV incidence data. The dates at which incidence in each of 151 districts in Southern Thailand exceeded specified thresholds were the target of model fits. We confronted multiple alternative models to determine which factors were most influential in the spatial spread. We considered multiple measures of spatial distance between districts and adjacency networks and also looked for evidence of long-distance translocation (LDT) events. The best fit model included driving-distance between districts, human movement, rubber plantation area and three LDT events. This work has important implications for predicting the spatial spread and targeting resources for control in future CHIKV emergences. Our modelling framework could also be adapted to other disease systems where population mobility may drive the spatial advance of outbreaks.

Behavioral responses to resource heterogeneity can accelerate biological invasions.

The abundance and spatial distribution of resources in a landscape and the behavioral response of individuals determines whether and how fast an invasive species spreads in an environment. Whether and how landscape manipulations can be used to slow invasive species is of great interest, in particular in forest ecosystems, where tree removal, thinning, and increasing tree diversity are discussed as management options. Classically, the focus is on availability and accessibility of resources; more recent considerations include individual-level behavioral movement responses to a spatially heterogeneous resource distribution. We derive a novel model for insect-host dynamics that includes three common behavioral aspects of foraging: higher movement rate in resource-poor areas, lower ovipositioning rate in resource-poor areas, and movement preference for resource-rich areas. We show that each of these basic mechanisms can increase the speed of invasion in a source-sink landscape above that in a homogeneous landscape with larger overall resource availability. We parameterize our model and illustrate our results with data for emerald ash borer, a recent highly destructive forest pest in North America. Our results highlight the importance of empirical work on movement behavior in different landscape types and near the interface between types.

Genetic mixture of multiple source populations accelerates invasive range expansion.

A wealth of population genetic studies have documented that many successful biological invasions stem from multiple introductions from genetically distinct source populations. Yet, mechanistic understanding of whether and how genetic mixture promotes invasiveness has lagged behind documentation that such mixture commonly occurs. We conducted a laboratory experiment to test the influence of genetic mixture on the velocity of invasive range expansion. The mechanistic basis for effects of genetic mixture could include evolutionary responses (mixed invasions may harbour greater genetic diversity and thus elevated evolutionary potential) and/or fitness advantages of between-population mating (heterosis). If driven by evolution, positive effects of source population mixture should increase through time, as selection sculpts genetic variation. If driven by heterosis, effects of mixture should peak following first reproductive contact and then dissipate. Using a laboratory model system (beetles spreading through artificial landscapes), we quantified the velocity of range expansion for invasions initiated with one, two, four or six genetic sources over six generations. Our experiment was designed to test predictions corresponding to the evolutionary and heterosis mechanisms, asking whether any effects of genetic mixture occurred in early or later generations of range expansion. We also quantified demography and dispersal for each experimental treatment, since any effects of mixture should be manifest in one or both of these traits. Over six generations, invasions with any amount of genetic mixture (two, four and six sources) spread farther than single-source invasions. Our data suggest that heterosis provided a 'catapult effect', leaving a lasting signature on range expansion even though the benefits of outcrossing were transient. Individual-level trait data indicated that genetic mixture had positive effects on local demography (reduced extinction risk and enhanced population growth) during the initial stages of invasion but no consistent effects on dispersal ability. Our work is the first to demonstrate that genetic mixture can alter the course of spatial expansion, the stage of invasion typically associated with the greatest ecological and economic impacts. We suggest that similar effects of genetic mixture may be a common feature of biological invasions in nature, but that these effects can easily go undetected.

The Influence of Evolution on Population Spread through Patchy Landscapes.

Predicting the spatial advance of biological invasions and range-shifting native species under climate change requires understanding how evolutionary processes influence the velocity of spread. Although landscape heterogeneity and the finite nature of individuals are known to influence the ecological dynamics of spreading populations, their effect on evolutionary dynamics influencing spread is poorly understood. We used adaptive dynamics and simulation approaches to evaluate the direction of selection on demographic and competitive traits in annual plant populations spreading through linear landscapes. In contrast to models in homogeneous environments where low-density fecundity is selectively favored, we found that large gaps between suitable habitat could favor the rise of competitively tolerant individuals at the invasion front, even when this ability to tolerate competitors came at the cost of reduced fecundity when rare. Simulations of the same processes incorporating finite individuals revealed a spatial priority effect; the long-term outcome of evolution strongly depended on which strategy initially got ahead. Finally, we found that although evolutionary change in demographic and competitive traits could increase the spread velocity, this increase could be insignificant in patchy landscapes, where competitive traits were favored in both spreading and nonspreading populations.

Spatial Transmission Characteristics of the Bluetongue Virus Serotype 3 Epidemic in The Netherlands, 2023.

A devastating bluetongue (BT) epidemic caused by bluetongue virus serotype 3 (BTV-3) has spread throughout most of the Netherlands within two months since the first infection was officially confirmed in the beginning of September 2023. The epidemic comes with unusually strong suffering of infected cattle through severe lameness, often resulting in mortality or euthanisation for welfare reasons. In total, tens of thousands of sheep have died or had to be euthanised. By October 2023, more than 2200 locations with ruminant livestock were officially identified to be infected with BTV-3, and additionally, ruminants from 1300 locations were showing BTV-associated clinical symptoms (but not laboratory-confirmed BT). Here, we report on the spatial spread and dynamics of this BT epidemic. More specifically, we characterized the distance-dependent intensity of the between-holding transmission by estimating the spatial transmission kernel and by comparing it to transmission kernels estimated earlier for BTV-8 transmission in Northwestern Europe in 2006 and 2007. The 2023 BTV-3 kernel parameters are in line with those of the transmission kernel estimated previously for the between-holding spread of BTV-8 in Europe in 2007. The 2023 BTV-3 transmission kernel has a long-distance spatial range (across tens of kilometres), evidencing that in addition to short-distance dispersal of infected midges, other transmission routes such as livestock transports probably played an important role.

High pathogenic avian influenza A(H5) viruses of clade 2.3.4.4b in Europe-Why trends of virus evolution are more difficult to predict.

Since 2016, A(H5Nx) high pathogenic avian influenza (HPAI) virus of clade 2.3.4.4b has become one of the most serious global threats not only to wild and domestic birds, but also to public health. In recent years, important changes in the ecology, epidemiology, and evolution of this virus have been reported, with an unprecedented global diffusion and variety of affected birds and mammalian species. After the two consecutive and devastating epidemic waves in Europe in 2020-2021 and 2021-2022, with the second one recognized as one of the largest epidemics recorded so far, this clade has begun to circulate endemically in European wild bird populations. This study used the complete genomes of 1,956 European HPAI A(H5Nx) viruses to investigate the virus evolution during this varying epidemiological outline. We investigated the spatiotemporal patterns of A(H5Nx) virus diffusion to/from and within Europe during the 2020-2021 and 2021-2022 epidemic waves, providing evidence of ongoing changes in transmission dynamics and disease epidemiology. We demonstrated the high genetic diversity of the circulating viruses, which have undergone frequent reassortment events, providing for the first time a complete overview and a proposed nomenclature of the multiple genotypes circulating in Europe in 2020-2022. We described the emergence of a new genotype with gull adapted genes, which offered the virus the opportunity to occupy new ecological niches, driving the disease endemicity in the European wild bird population. The high propensity of the virus for reassortment, its jumps to a progressively wider number of host species, including mammals, and the rapid acquisition of adaptive mutations make the trend of virus evolution and spread difficult to predict in this unfailing evolving scenario.

Evolution of dispersal and life history interact to drive accelerating spread of an invasive species.

Populations on the edge of an expanding range are subject to unique evolutionary pressures acting on their life-history and dispersal traits. Empirical evidence and theory suggest that traits there can evolve rapidly enough to interact with ecological dynamics, potentially giving rise to accelerating spread. Nevertheless, which of several evolutionary mechanisms drive this interaction between evolution and spread remains an open question. We propose an integrated theoretical framework for partitioning the contributions of different evolutionary mechanisms to accelerating spread, and we apply this model to invasive cane toads in northern Australia. In doing so, we identify a previously unrecognised evolutionary process that involves an interaction between life-history and dispersal evolution during range shift. In roughly equal parts, life-history evolution, dispersal evolution and their interaction led to a doubling of distance spread by cane toads in our model, highlighting the potential importance of multiple evolutionary processes in the dynamics of range expansion.

Density-dependent within-patch movement behavior of two competing species.

Movement behavior is central to understanding species distributions, population dynamics and coexistence with other species. Although the relationship between conspecific density and emigration has been well studied, little attention has been paid to how interspecific competitor density affects another species' movement behavior. We conducted releases of two species of competing Tribolium flour beetles at different densities, alone and together in homogeneous microcosms, and tested whether their recaptures-with-distance were well described by a random-diffusion model. We also determined whether mean displacement distances varied with the release density of conspecific and heterospecific beetles. A diffusion model provided a good fit to the redistribution of T. castaneum and T. confusum at all release densities, explaining an average of >60% of the variation in recaptures. For both species, mean displacement (directly proportional to the diffusion rate) exhibited a humped-shaped relationship with conspecific density. Finally, we found that both species of beetle impacted the within-patch movement rates of the other species, but the effect depended on density. For T. castaneum in the highest density treatment, the addition of equal numbers of T. castaneum or T. confusum had the same effect, with mean displacements reduced by approximately one half. The same result occurred for T. confusum released at an intermediate density. In both cases, it was total beetle abundance, not species identity that mattered to mean displacement. We suggest that displacement or diffusion rates that exhibit a nonlinear relationship with density or depend on the presence or abundance of interacting species should be considered when attempting to predict the spatial spread of populations or scaling up to heterogeneous landscapes.

Science paper or big data? Assessing invasion dynamics using observational data.

Non-native species are spreading at an unprecedented rate over large spatial scales, with global environmental change and growth in commerce providing novel opportunities for range expansion. Assessing the pattern and rate of spread is key to the development of strategies for safeguarding against future invasions and efficiently managing existing ones. Such assessments often depend on spatial distribution data from online repositories, which can be spatially biased, imprecise, and lacking in quantity. Here, the influence of disparities between occurrence records from online data repositories and what is known of the invasion history from peer-reviewed published literature on non-native species range expansion was evaluated using 6693 records of the Pacific oyster, Magallana gigas (Thunberg, 1793), spanning 56 years of its invasion in Europe. Two measures of spread were calculated: maximum rate of spread (distance from introduction site over time) and accumulated area (spatial expansion). Results suggest that despite discrepancies between online and peer-reviewed data sources, including a paucity of records from the early invasion history in online repositories, the use of either source does not result in significantly different estimates of spread. Our study significantly improves our understanding of the European distribution of M. gigas and suggests that a combination of short- and long-range dispersal drives range expansions. More widely, our approach provides a framework for comparison of online occurrence records and invasion histories as documented in the peer-reviewed literature, allowing critical evaluation of both data sources and improving our understanding of invasion dynamics significantly.

Relative timing of respiratory syncytial virus epidemics in summer 2021 across the United States was similar to a typical winter season.

We used a validated proxy of respiratory syncytial virus (RSV) activity in the United States (Google search data) to evaluate the onsets of RSV epidemics in 2021 and 2016-2019. Despite the unusual out-of-season summer timing, the relative timing of RSV epidemics between states in 2021 shared a similar spatial pattern with typical winter RSV seasons. Our results suggest that the onset of RSV epidemics in Florida can serve as a baseline to adjust the initiation of prophylaxis administration and clinical trials in other states regardless of the seasonality of RSV epidemics.

HCV Spread Kinetics Reveal Varying Contributions of Transmission Modes to Infection Dynamics.

The hepatitis C virus (HCV) is capable of spreading within a host by two different transmission modes: cell-free and cell-to-cell. However, the contribution of each of these transmission mechanisms to HCV spread is unknown. To dissect the contribution of these different transmission modes to HCV spread, we measured HCV lifecycle kinetics and used an in vitro spread assay to monitor HCV spread kinetics after a low multiplicity of infection in the absence and presence of a neutralizing antibody that blocks cell-free spread. By analyzing these data with a spatially explicit mathematical model that describes viral spread on a single-cell level, we quantified the contribution of cell-free, and cell-to-cell spread to the overall infection dynamics and show that both transmission modes act synergistically to enhance the spread of infection. Thus, the simultaneous occurrence of both transmission modes represents an advantage for HCV that may contribute to viral persistence. Notably, the relative contribution of each viral transmission mode appeared to vary dependent on different experimental conditions and suggests that viral spread is optimized according to the environment. Together, our analyses provide insight into the spread dynamics of HCV and reveal how different transmission modes impact each other.

Phenomenological model of auditory nerve population responses to cochlear implant stimulation.

BACKGROUND: Models of auditory nerve fiber (ANF) responses to electrical stimulation are helpful to develop advanced coding for cochlear implants (CIs). A phenomenological model of ANF population responses to CI electrical stimulation with a lower computational complexity compared to a biophysical model would be beneficial to evaluate new CI coding strategies. NEW METHOD: This study presents a phenomenological model which combines four temporal characteristics of ANFs (refractoriness, facilitation, accommodation and spike rate adaptation) in addition to a spatial spread of the electric field. RESULTS: The model predicts the performances of CI subjects in the melodic contour identification (MCI) experiment. The simulations for the MCI experiment were consistent with CI recipients' experimental outcomes that were not predictable from the electrical stimulation patterns themselves. COMPARISON WITH EXISTING METHODS: Previously, no phenomenological population model of ANFs has combined all four aforementioned temporal phenomena. CONCLUSIONS: The proposed model would help the further investigations of ANFs responses to different electrical stimulation patterns and comparison of different sound coding strategies in CIs.

Spatial spread and emergence of reassortant H5 highly pathogenic avian influenza viruses in Iran.

Since 2005, H5Nx highly pathogenic avian influenza (HPAI) viruses of the Goose/Guangdong (Gs/GD) lineage have spread worldwide, affecting poultry and wild birds in Asia, Europe, Africa and North America. So far, the role of Western Asia and the Middle East in the diffusion dynamics of this virus has been poorly explored. In order to investigate the genetic diversity and the role of Iran in the transmission dynamics of the Gs/GD lineage, we sequenced the complete genome of twenty-eight H5Nx viruses which were circulating in the country between 2016 and 2018. We reported the first characterization of the HPAI H5N6 subtype of clade 2.3.4.4B in Iran and gave evidence of the high propensity of the Gs/GD H5 AIVs to reassort, describing six novel H5N8 genotypes of clade 2.3.4.4B, some of them likely generated in this area, and one H5N1 reassortant virus of clade 2.3.2.1c. Our spatial analyses demonstrated that the viruses resulted from different viral introductions from Asia and Europe and provided evidence of virus spread from Iran to the Middle East. Therefore, Iran may represent a hot-spot for virus introduction, dissemination and for the generation of new genetic variability. Increasing surveillance efforts in this high-risk area is of utmost importance for the early detection of novel emerging strains with zoonotic potential.

Loss of cytoplasmic incompatibility and minimal fecundity effects explain relatively low Wolbachia frequencies in Drosophila mauritiana.

Maternally transmitted Wolbachia bacteria infect about half of all insect species. Many Wolbachia cause cytoplasmic incompatibility (CI) and reduced egg hatch when uninfected females mate with infected males. Although CI produces a frequency-dependent fitness advantage that leads to high equilibrium Wolbachia frequencies, it does not aid Wolbachia spread from low frequencies. Indeed, the fitness advantages that produce initial Wolbachia spread and maintain non-CI Wolbachia remain elusive. wMau Wolbachia infecting Drosophila mauritiana do not cause CI, despite being very similar to CI-causing wNo from Drosophila simulans (0.068% sequence divergence over 682,494 bp), suggesting recent CI loss. Using draft wMau genomes, we identify a deletion in a CI-associated gene, consistent with theory predicting that selection within host lineages does not act to increase or maintain CI. In the laboratory, wMau shows near-perfect maternal transmission; but we find no significant effect on host fecundity, in contrast to published data. Intermediate wMau frequencies on the island of Mauritius are consistent with a balance between unidentified small, positive fitness effects and imperfect maternal transmission. Our phylogenomic analyses suggest that group-B Wolbachia, including wMau and wPip, diverged from group-A Wolbachia, such as wMel and wRi, 6-46 million years ago, more recently than previously estimated.