THE EFFECTS OF MATING SYSTEM AND GENETIC VARIABILITY ON SUSCEPTIBILITY TO TREMATODE PARASITES IN A FRESHWATER SNAIL, LYMNAEA STAGNALIS

The amount and distribution of genetic variability in host populations can have significant effects on the outcome of host-parasite interactions. We studied the effect of mating system and genetic variability on susceptibility of Lymnaea stagnalis snails to trematode parasites. Mating system of snails from eight populations differing in the amount of genetic variability was manipulated, and self- and cross-fertilized offspring were exposed to naturally occurring trematode parasites in a controlled lake experiment. Susceptibility of snails varied between populations, but mating-system treatment did not have a significant effect. Heterozygosity of snails was negatively correlated with the pro…

Coexistence and resource competition.

How large numbers of species coexist on a seemingly limited number of different resources is a classic problem in ecology1, and attempts have been made to solve it experimentally. But we are not convinced that Huisman and Weissing's2 proposal to add non-stationary dynamics in species abundance to the list of possible explanations offers any new insight into this biodiversity enigma.

Methane production and oxidation potentials in relation to water table fluctuations in two boreal mires

We studied the response of methane production and oxidation potentials in a minerotrophic and an ombrotrophic mire to water table fluctuations. In profiles where water table had not varied, the water-saturated layers showed significant potentials while the unsaturated layers did not. The production potentials in the saturated layers below water level ranged from 0.1 to 2.4 m gC H4 h ˇ1 (g d.w.) ˇ 1 and oxidation potentials (first order reaction rate constants) betweenˇ0.010 andˇ0.120 h ˇ1 (g d.w.) ˇ 1 . In profiles with constant water level, the maximal production potential occurred 20 cm and maximal oxidation potential 10 cm below water level. When water table varied only a little, product…

Species loss leads to community closure

Global extinction of a species is sadly irreversible. At a local scale, however, extinctions may be followed by re-invasion. We here show that this is not necessarily the case and that an ecological community may close its doors for re-invasion of species lost from it. Previous studies of how communities are assembled have shown that there may be rules for that process and that limitations are set to the order by which species are introduced and put together. Instead of focusing on the assembly process we randomly generated simple competitive model communities that were stable and allowed for two to 10 coexisting species. When a randomly selected single species was removed from the communit…

A mechanistic underpinning for sigmoid dose-dependent infection

Self-organized dynamics in spatially structured populations

Self-organization and pattern formation represent the emergence of order in temporal and spatial processes. Self-organization in population ecology is gaining attention due to the recent advances concerning temporal fluctuations in the population size of dispersal-linked subunits. We shall report that spatially structured models of population renewal promote the emergence of a complex power law order in spatial population dynamics. We analyse a variety of population models showing that self-organization can be identified as a temporal match in population dynamics among local units, and how the synchrony changes in time. Our theoretical results are concordant with analyses of population data…

Harvesting‐induced population fluctuations?

It has recently been shown that damped endogenous dynamics is a common feature in Finnish grouse species; In this paper, we demonstrate that time-variant harvesting may turn damped dynamics to quasi-periodic fluctuations. Exploited populations, e.g. grouse, may therefore fluctuate more than expected if we do not manage to keep the harvest fraction constant over time. However, the harvest fraction of Finnish grouse varies with the phase of the cycle. Such a harvesting strategy could potentially change the periodicity of the fluctuations, as can a threshold harvest strategy where a constant fraction is harvested above a density threshold. The two non-linear harvesting strategies investigated …

Primers for nine microsatellite loci in the hermaphroditic snailLymnaea stagnalis

Variation in and amplification conditions for nine polymorphic microsatellite loci identified from Lymnaea stagnalis , a hermaphroditic pulmonate snail, are described. Eight populations from central Finland were studied, which varied in terms of both observed polymorphism and heterozygosity. The number of alleles at each locus is moderate (two to seven), except for one exceptional locus having 16 alleles, and for which null alleles are possible. There is no evidence for genotypic disequilibrium in the populations for all pairs of loci. Heterozygosity levels are indicative of outcrossing in L. stagnalis , whose mating system will be characterized further using these markers.

Interactions between ecological traits and host plant type explain distribution change in noctuid moths.

The ecological traits of species determine how well a species can withstand threats to which it is exposed. If these predisposing traits can be identified, species that are most at risk of decline can be identified and an understanding of the processes behind the declines can be gained. We sought to determine how body size, specificity of larval host plant, overwintering stage, type of host plant, and the interactions of these traits are related to the distribution change in noctuid moths. We used data derived from the literature and analyzed the effects of traits both separately and simultaneously in the same model. When we analyzed the traits separately, it seemed the most important deter…

Community structure and the evolution of aposematic coloration

Studies on the evolution of aposematic coloration (prey coloration advertising for unpalatability) have mainly focused on predator psychology in simplified single-prey species systems. We chose, instead, to model population dynamics on the community level. We studied the invasion by an aposematic phenotype in the presence and absence of another prey species. The single-prey and two-prey models differed in two major ways. First, with two prey species the invasion was possible only with a weak aposematic signal, whereas with a single prey species there was no such an upper limit for signal strength. Second, with a single prey species, increase of the aposematic phenotype always resulted in ra…

Environmental noise and population dynamics of the ciliated protozoa Tetrahymena thermophila in aquatic microcosms

Population theory predicts that the reddened environmental noise, especially in combination with high population growth rate, reddens population dynamics, increases population variability and strengthens environment–population correlation. We tested these predictions with axenic populations of ciliated protozoa Tetrahymena thermophila. Populations with low and high growth rate were cultured in a stable environment, and in environments with sublethal temperature fluctuations that had blue, white and red spectra (i.e. negatively autocorrelated, uncorrelated, or positively autocorrelated, respectively). Population size and biomass of individuals were determined at 3-h intervals for 18 days. Dy…

Sex in space: population dynamic consequences

Sex, so important in the reproduction of bigametic species, is nonetheless often ignored in explorations of the dynamics of populations. Using a growth model of dispersal-coupled populations we can keep track of fluctuations in numbers of females and males. The sexes may differ from each other in their ability to disperse and their sensitivity to population density. As a further complication, the breeding system is either monogamous or polygamous. We use the harmonic mean birth function to account for sex-ratio-dependent population growth in a Moran–Ricker population renewal process. Incorporating the spatial dimension stabilizes the dynamics of populations with monogamy as the breeding sys…

Interactions between environmental variability and immigration rate control patterns of species diversity

Abstract Theories focussing on local competition processes predict that species diversity is maximised with high-frequency environmental variability or alternatively with intermediate frequencies. The models coupling regional processes to local explanations of diversity patterns predict that the immigration rate from a regional species pool can strongly increase diversity. However, the interaction between local and regional processes in temporally fluctuating environment has received little attention. We explored in a simulation study how the patterns of species diversity are affected by the frequency spectrum of stochastic environmental variations and density independent immigration rate i…

Evolutionary population dynamics

The interface between the evolution of life history traits and population dynamics in temporally and spatially variable environments is the topic of this chapter. Thus, the frame for the life history processes is set by spatial and temporal fluctuations in population density. Here, we will focus primarily on modes of reproduction and we are especially interested in whether alternative reproductive strategies can co-exist in a population. We show that spatially structured populations may allow co-existence of various life history strategies that do not easily co-exist in a nonstructured environment. Also, intrinsic and external temporal fluctuations in the environment tend to enhance polymor…

Resource availability and goshawk offspring sex ratio variation: a large-scale ecological phenomenon

Summary 1. Local population studies have shown that sex allocation among many birds and mammals seems to be partly non-random and in connection to surrounding factors, such as environmental or parental quality. In this scenario, if environmental quality varies in space and time, it is feasible that environmental quality also comes to influence offspring sex ratio on larger geographical scales. 2. Investigating this idea - using nation-wide data sets on size-dimorphic Finnish northern goshawks Accipiter gentilis from 1989 to 1998 - we found that offspring sex ratio is related to spatial and temporal variation in availability of their main prey, woodland grouse species. 3. In a majority of lo…

Seed Bank in Annuals: Competition Between Banker and Non-banker Morphs

Seed bank is a plant life history strategy against the unpredictability of the biotic and the abiotic environment. We simulated competition between a seed banking and a non-banking morph of an annual plant. A constant fraction of the banker morph seeds was allocated to the seed bank, where they had a constant mortality and germination rate. All surviving seeds of the non-banker morph germinated in the next generation. The seedlings of both morphs experienced similar density-dependent mortality. Whether one of the morphs wins or the morphs coexist was evaluated from parameter space plots and statistically with logistic regression analysis. All parameters of the model had a significant, nonli…

Mate‐Search Efficiency Can Determine the Evolution of Separate Sexes and the Stability of Hermaphroditism in Animals

Limited availability of mating partners has been proposed as an explanation for the occurrence of simultaneous hermaphroditism in animals with pair mating. When low population density or low mobility of a species limits the number of potential mates, simultaneous hermaphrodites may have a selective advantage because, first, they are able to adjust the allocation of resources between male and female functions in order to maximize fitness; second, in a hermaphroditic population the likelihood of meeting a partner is higher because all individuals are potential mates; and, third, in the absence of mating partners, many simultaneously hermaphroditic animals have the option of reproducing throug…

The Spatial Dimension in Population Fluctuations

Theoretical research into the dynamics of coupled populations has suggested a rich ensemble of spatial structures that are created and maintained either by external disturbances or self-reinforcing interactions among the populations. Long-term data of the Canadian lynx from eight Canadian provinces display large-scale spatial synchrony in population fluctuations. The synchronous dynamics are not time-invariant, however, as pairs of populations that are initially in step may drift out of phase and back into phase. These observations are in agreement with predictions of a spatially-linked population model and support contemporary population ecology theory.

Non-linear biological responses to disturbance: consequences on population dynamics

Abstract We assessed how non-linear biological responses to environmental noise, or “noise filtering”, impact the spectra of density-dependent population dynamics, and the correlation between noise and population dynamics. The noise was assumed to affect population growth rate in a discrete-time population model by Hassell [J. Anim. Ecol. 44 (1975) 283–295] where the population growth rate was linked to the environment with an optimum type filter. When compared to unfiltered noise, the filtered noise can distort the stationary distribution of population values. The optimum type filter can make cyclic population dynamics more regular and low population values can become more frequent or rare…

SHARING THE BENEFITS OF COOPERATION IN HIGH SEAS FISHERIES: A CHARACTERISTIC FUNCTION GAME APPROACH

Temporal variability in detritus resource maintains diversity of bacterial communities

Competition theory generally predicts that diversity is maintained by temporal environmental fluctuations. One of the many suggested mechanisms for maintaining diversity in fluctuating environments is the gleaner-opportunist trade-off, whereby gleaner species have low threshold resource levels and low maximum growth rates in high resource concentration while opportunist species show opposite characteristics. We measured the growth rates of eight heterotrophic aquatic bacteria under different concentrations of chemically complex plant detritus resource. The growth rates revealed gleaner-opportunist trade-offs. The role of environmental variability in maintaining diversity was tested in a 28-…

Punishment of polygyny

We investigated the evolution of monogamy (one male, one female) and polygyny (one male, more than one female). In particular, we studied whether it is possible for a mutant polygynous mating strategy to invade a resident population of monogamous breeders and, alternatively, whether a mutant monogamy can invade resident polygyny. Our population obeys discrete-time Ricker dynamics. The role of males and females in the breeding system is incorporated via the harmonic birth function. The results of the invasability analysis are straightforward. Polygyny is an evolutionarily stable strategy mating system; this holds throughout the examined range of numbers of offspring produced per female. So t…

Ecological correlates of distribution change and range shift in butterflies

1. In order to be effective custodians of biodiversity, one must understand what ecological characteristics predispose species to population decline, range contraction, and, eventually, to extinction. 2. The present paper analyses distribution change (area of occupancy) and range shift (extent and direction) of the threatened and non-threatened butterfly species in Finland, and identifies species-specific ecological characteristics promoting changes in distribution and range. 3. Overall, the range of butterflies has shifted along the climatic isotherms, suggesting that climate change has influenced species’ ranges. Interestingly, though, threatened species have moved very little and not to …

From arctic lemmings to adaptive dynamics: Charles Elton's legacy in population ecology.

We shall examine the impact of Charles S. Elton's 1924 article on periodic fluctuations in animal populations on the development of modern population ecology. We argue that his impact has been substantial and that during the past 75 years of research on multi-annual periodic fluctuations in numbers of voles, lemmings, hares, lynx and game animals he has contributed much to the contemporary understanding of the causes and consequences of population regulation. Elton was convinced that the cause of the regular fluctuations was climatic variation. To support this conclusion, he examined long-term population data then available. Despite his firm belief in a climatic cause of the self-repeating …

Environmental Variability and Semelparity vs. Iteroparity as Life Histories

Research on the evolution of life histories addresses the topic of fitness trade-offs between semelparity (reproducing once in a lifetime) and iteroparity (repeated reproductive bouts per lifetime). Bulmer (1994) derived the relationship v+P(A)<1 (P(A) is the adult survival;vb(S) and b(S) are the offspring numbers for iteroparous and semelparous breeding strategies, respectively), under which a resident semelparous population cannot be invaded by an iteroparous mutant when the underlying population dynamics are stable. We took Bulmer's population dynamics, and added noise in juvenile and adult survival and in offspring numbers. Long-term coexistence of the two strategies is possible in much…

Predation and resource fluctuations drive eco-evolutionary dynamics of a bacterial community

Predation and temporal resource availability are among the most important factors determining prey community dynamics and composition. Both factors have been shown to affect prey diversity, but less is known about their interactive effects, especially in rapidly evolving prey communities. In a laboratory microcosm experiment, we manipulated the presence of the predatory protozoan Tetrahymena thermophila and the temporal patterns in the availability of resources for a bacterial prey community. We found that both predation and temporal fluctuations in prey resources resulted in a more even prey community, and these factors also interacted so that the effect of predation was only seen in a flu…

Is the impact of environmental noise visible in the dynamics of age-structured populations?

Climate change has ignited lively research into its impact on various population–level processes. The research agenda in ecology says that some of the fluctuations in population size are accountable for by the external noise (e.g. weather) modulating the dynamics of populations. We obeyed the agenda by assuming population growth after a resource–limited Leslie matrix model in an age–structured population. The renewal process was disturbed by superimposing noise on the development of numbers in one or several age groups. We constructed models for iteroparous and semelparous breeders so that, for both categories, the population growth rate was matching. We analysed how the modulated populatio…

Ecological Determinants of Distribution Decline and Risk of Extinction in Moths

For successful conservation of species it is important to identify traits that predispose species to the risk of extinction. By identifying such traits conservation efforts can be directed toward species that are most at risk of becoming threatened. We used data derived from the literature to determine ecological traits that affect distribution, distribution change, and the risk of extinction in Finnish noctuid moths (Lepidoptera, Noctuidae). The ecological traits we examined included body size, larval specificity, length of the flight period, and overwintering stage. In addition, in monophagous species we examined the effects of resource distribution. Larval specificity, length of the flig…

Conditions for the spread of conspicuous warning signals: A numerical model with novel insights

The initial evolution of conspicuous warning signals presents an evolutionary problem because selection against rare conspicuous signals is presumed to be strong, and new signals are rare when they first arise. Several possible solutions have been offered to solve this apparent evolutionary paradox, but disagreement persists over the plausibility of some of the proposed mechanisms. In this paper, we construct a deterministic numerical simulation model that allows us to derive the strength of selection on novel warning signals in a wide range of biologically relevant situations. We study the effects of predator psychology (learning, rate of mistaken attacks, and neophobia) on selection. We a…

Species immigration, extinction and turnover of vascular plants in boreal lakes

Dictated by limited resource availability for land acquisition, a central question in conservation biology is the ability of areas of different size to maintain species diversity. The selected reserves should not only be species rich at the moment, but should also maintain species diversity in the long run. We used two sets of data on vascular plant species in boreal lakes collected in 1933/34 and 1996 to test the relationships between lake area and the extinction, immigration and turnover rates of the species. Moreover, we investigated, whether the number of species in 1933/34 or water connection between lakes was related to extinction, immigration and turnover rates of species. We found t…

The irreducible uncertainty of the demography–environment interaction in ecology

The interpretation of ecological data has been greatly improved by bridging the gap between ecological and statistical models. The major challenge is to separate competing hypotheses concerning demography, or other ecological relationships, and environmental variability (noise). In this paper we demonstrate that this may be an arduous, if not impossible, task. It is the lack of adequate ecological theory, rather than statistical sophistication, which leads to this problem. A reconstruction of underlying ecological processes can only be done if we are certain of either the demographic or the noise model, which is something that can only be achieved by an improved theory of stochastic ecologi…

EVOLUTION OF HSP90 EXPRESSION IN TETRAHYMENA THERMOPHILA (PROTOZOA, CILIATA) POPULATIONS EXPOSED TO THERMALLY VARIABLE ENVIRONMENTS

Evolutionary consequences of thermally varying environments were studied in the ciliated protozoan Tetrahymena thermophila. Replicated lines were propagated for 60 days, a maximum of 500 generations, in stable, slowly fluctuating (red spectrum), and rapidly fluctuating (blue spectrum) temperatures. The red and blue fluctuations had a dominant period length of 15 days and two hours, respectively. The mean temperature of all time series was 25 degrees C and the fluctuating temperatures had the same minimum (10 degrees C), maximum (40 degrees C), and variance. During the experiment, population sizes and biomasses were monitored at three-day intervals. After the experiment, carrying capacity an…

Maternal effects and the stability of population dynamics in noisy environments

Summary 1. It is widely appreciated that complex population dynamics are more likely in systems where there is a lag in the density dependence. The transmission of maternal environmental conditions to offspring phenotype is a potential cause of such a lag. Maternal effects are increasingly found to be common in a wide range of organisms, and might thus be a frequent cause of nonequilibrium population dynamics. 2. We show that a maternal effects’ lag generally increases population variability. This may result from the lag inducing cycles (or more complex dynamics) in a deterministic environment or, in a stochastic environment, from the lag interacting with environmental noise to produce more…

Population dynamic consequences of delayed life-history effects

Evidence from wildlife and human populations indicates that conditions during early development can have marked effects on the subsequent performance of individuals and cohorts. Likewise, the effects of maternal and, more generally, parental environments can be transferred among individuals between generations. These delayed life-history effects are found consistently and suggestions have been made that they can be one source of both variability and of delayed density dependence in population dynamics. Assessments of several different time series indicate that population variability and delayed density dependence are common and that understanding the mechanisms giving rise to them is crucia…

Invasion Ability and Disease Dynamics of Environmentally Growing Opportunistic Pathogens under Outside-Host Competition

Most theories of the evolution of virulence concentrate on obligatory host-pathogen relationship. Yet, many pathogens replicate in the environment outside-host where they compete with non-pathogenic forms. Thus, replication and competition in the outside-host environment may have profound influence on the evolution of virulence and disease dynamics. These environmentally growing opportunistic pathogens are also a logical step towards obligatory pathogenicity. Efficient treatment methods against these diseases, such as columnaris disease in fishes, are lacking because of their opportunist nature. We present a novel epidemiological model in which replication and competition in the outside-hos…

The use of ecological traits in extinction risk assessments: A case study on geometrid moths

Identifying ecological traits that make some species more vulnerable than others is vital for predictive conservation science. By identifying these predisposing traits we can predict which species are most prone to decline and gain an understanding of the reasons behind the decline. The aim of this study was to determine the ecological traits that best predict extinction risk and distribution change in Finnish geometrid moths and to develop an understanding of the biological connections between these traits and threats. We found that larval specificity, overwintering stage and flight period length predicted distribution change and extinction risk. There was also an interaction effect betwee…

Population variability in space and time.

One of the most ubiquitous phenomena of all natural populations is their variability in numbers in space and time. However, there are notable differences among populations in the way the population size fluctuates. One of the major challenges in population and community ecology is to explain and understand this variety and to find possible underlying rules that might be modified from case-to-case. Population variability also has a spatial component because fluctuations are often synchronized over relatively large distances. Recently, this has led to growing interest in how 'internal' (density-dependent) processes interact with 'external' factors such as environmental variability.

Does evolution of iteroparous and semelparous reproduction call for spatially structured systems?

A persistent question in the evolution of life histories is the fitness trade-off between reproducing only once (semelparity) in a lifetime or reproducing repeated times in different seasons (iteroparity). The problem can be formulated into a research agenda by assuming that one reproductive strategy is resident (has already evolved) and by asking whether invasion (evolution) of an alternative reproductive strategy is possible. For a spatially nonstructured system, Bulmer (1994) derived the relationship v + PA1 (PA is adult survival; vbS and bS are offspring numbers for iteroparous and semelparous breeding strategies, respectively) at which semelparous population cannot be invaded by an ite…

Non-unique population dynamics: basic patterns

We review the basic patterns of complex non-uniqueness in simple discrete-time population dynamics models. We begin by studying a population dynamics model of a single species with a two-stage, two-habitat life cycle. We then explore in greater detail two ecological models describing host‐macroparasite and host‐parasitoid interspecific interactions. In general, several types of attractors, e.g. point equilibria vs. chaotic, periodic vs. quasiperiodic and quasiperiodic vs. chaotic attractors, may coexist in the same mapping. This non-uniqueness also indicates that the bifurcation diagrams, or the routes to chaos, depend on initial conditions and are therefore non-unique. The basins of attrac…

GENETIC VARIABILITY AND DRIFT LOAD IN POPULATIONS OF AN AQUATIC SNAIL

Population genetic theory predicts that in small populations, random genetic drift will fix and accumulate slightly deleterious mutations, resulting in reduced reproductive output. This genetic load due to random drift (i.e., drift load) can increase the extinction risk of small populations. We studied the relationship between genetic variability (indicator of past population size) and reproductive output in eight isolated, natural populations of the hermaphroditic snail Lymnaea stagnalis. In a common laboratory environment, snails from populations with the lowest genetic variability mature slower and have lower fecundity than snails from genetically more variable populations. This result s…

Dynamic complexities in host-parasitoid interaction

In the 1970s ecological research detected chaos and other forms of complex dynamics in simple population dynamics models, initiating a new research tradition in ecology. However, the investigations of complex population dynamics have mainly concentrated on single populations and not on higher dimensional ecological systems. Here we report a detailed study of the complicated dynamics occurring in a basic discrete-time model of host-parasitoid interaction. The complexities include (a) non-unique dynamics, meaning that several attractors coexist, (b) basins of attraction (defined as the set of the initial conditions leading to a certain type of an attractor) with fractal properties (pattern of…

Extinction risk under coloured environmental noise

Positively autocorrelated red environmental noise is characterized by a strong dependence of expected sample variance on sample length. This dependence has to be taken into account when assessing extinction risk under red and white uncorrelated environmental noise. To facilitate a comparison between red and white noise, their expected variances can be scaled to be equal, but only at a chosen time scale. We show with a simple one-dimensional population dynamics model that the different but equally reasonable choices of the time scale yield qualitatively different results on the dependence of extinction risk on the colour of environmental noise: extinction risk might increase as well as decre…

Predominance of outcrossing in Lymnaea stagnalis despite low apparent fitness costs of self-fertilization.

We have quantified the natural mating system in eight populations of the simultaneously hermaphroditic aquatic snail Lymnaea stagnalis, and studied the ecological and genetic forces that may be directing mating system evolution in this species. We investigated whether the natural mating system can be explained by the availability of mates, by the differential survival of self- and cross-fertilized snails in nature, and by the effects of mating system on parental fecundity and early survival. The natural mating system of L. stagnalis was found to be predominantly cross-fertilizing. Density of snails in the populations had no relationship with the mating system, suggesting that outcrossing ra…

A Tale of Big Game and Small Bugs

Animal populations undergo repetitive cycles of rising and falling numbers. In a Perspective, [Ranta and colleagues][1] discuss the value of time-series analyses for examining the changing dynamics of animal populations. A large time series gathered by the Hudson Bay Company based on the fur trade has provided invaluable data on the rise and fall of the Canadian lynx population from 1821 to the present. Analysis of this time series reveals that climate as well as factors influencing birth and death rates are important in regulating the lynx population ([ Stenseth et al .][2]). In a separate study, mathematical modeling combined with fieldwork revealed the importance of predators in determin…

Synchronicity in population systems: cause and consequence mixed

The enigma of frequency-dependent selection

Frequency-dependent selection is so fundamental to modern evolutionary thinking that everyone interested in evolutionary biology 'knows' the concept. It is even so fundamental that many authors of textbooks do not bother to define it. Yet it turns out that different authors (and sometimes even one and the same author) use the term to refer to different types of selection. In this paper we try to uncover the sources of this confusion. The concept is fairly well defined in the original concept of population genetical theory, which focuses on short-term evolutionary change, and basically ignores density-dependence. The problems start when the original concept is used in the context of long-ter…

How does environmental variation translate into biological processes?

Birth and death rates, as so many other biological processes, are usually not linearly related to environmental variation. Common examples of non-linear response forms include unimodal ‘‘optimum-type’’ responses and various saturating responses. These responses filter the signal coming from the environment to a corresponding biological process. We explored how different types of environmental signal may be transformed to a biological process. We were interested in the effect of the filter on modulation of (1) the variance of the signal, on (2) the variance-covariance structure between the signal and the filtered signal, and on (3) the match between the power spectra of the signal and the fi…

Reply from m. Heino, j.a.j. Metz and v. Kaitala.

Eva Kisdi clarifies the relationships between frequency dependence, optimization and ESSs. We basically agree with all her comments. However, some further clarification may be useful.In the first paragraph of Kisdi's letter, ESSs and optimal strategies are seemingly opposed by saying that `finding an optimal strategy is a considerably stronger result than finding an ESS'. Although this statement is factually correct, it might engender a suggestion that is slightly wrong. Conceptually, ESSs are always primary: only ESSs matter from the viewpoint of long-term evolution. Optimization is secondary only, and must be justified by an ESS argument that explicitly accounts for the ecology in which t…

Outside-host growth of pathogens attenuates epidemiological outbreaks.

Opportunist saprotrophic pathogens differ from obligatory pathogens due to their capability in host-independent growth in environmental reservoirs. Thus, the outside-host environment potentially influences host-pathogen dynamics. Despite the socio-economical importance of these pathogens, theory on their dynamics is practically missing. We analyzed a novel epidemiological model that couples outside-host density-dependent growth to host-pathogen dynamics. Parameterization was based on columnaris disease, a major hazard in fresh water fish farms caused by saprotrophic Flavobacterium columnare. Stability analysis and numerical simulations revealed that the outside-host growth maintains high pr…

Outside-host phage therapy as a biological control against environmental infectious diseases

Background Environmentally growing pathogens present an increasing threat for human health, wildlife and food production. Treating the hosts with antibiotics or parasitic bacteriophages fail to eliminate diseases that grow also in the outside-host environment. However, bacteriophages could be utilized to suppress the pathogen population sizes in the outside-host environment in order to prevent disease outbreaks. Here, we introduce a novel epidemiological model to assess how the phage infections of the bacterial pathogens affect epidemiological dynamics of the environmentally growing pathogens. We assess whether the phage therapy in the outside-host environment could be utilized as a biologi…

Predicting the risk of extinction from shared ecological characteristics

Understanding the ultimate causes of population declines and extinction is vital in our quest to stop the currently rampant biodiversity loss. Comparison of ecological characteristics between threatened and nonthreatened species may reveal these ultimate causes. Here, we report an analysis of ecological characteristics of 23 threatened and 72 nonthreatened butterfly species. Our analysis reveals that threatened butterflies are characterized by narrow niche breadth, restricted resource distribution, poor dispersal ability, and short flight period. Based on the characteristics, we constructed an ecological extinction risk rank and predicted which of the currently nonthreatened species are at …

Additional file 2: of Outside-host phage therapy as a biological control against environmental infectious diseases

Equilibrium population densities and the Jacobian matrices for phage therapy model (1)â (6). Local stability analysis of the P-S-I submodel at S=Iâ =â 0 in the absence of recovery. (DOCX 30 kb)

Outside-host predation as a biological control against an environmental opportunist disease

Abstract Environmentally growing opportunist pathogens are a common threat to human health and food production. Due to environmental growth of the pathogen, these diseases are difficult to control with disinfectants and antibiotics. Thus, there is a need for sustainable and effective control methods against environmentally growing opportunist diseases. Predation is often a major limiting factor in the outside host environment. Here we propose that it could be used in the biological control of these diseases. We introduce a novel epidemiological model for environmentally growing opportunists combining pathogen growth within-host (SI model) and outside-host into classical predator-prey model.…

Geographical patterns of species turnover in aquatic plant communities

1. A classic theory in biogeography predicts that high latitude communities are unstable. This may be because of decreased species richness or decreased environmental predictability and productivity towards the poles. 2. We studied latitudinal patterns in long-term community persistence of aquatic vascular plants in 112 Finnish lakes, situated within a 1000-km range from the northernmost to the southernmost lake. 3. Contrary to theoretical predictions, we found that the turnover rate of plant species in 45 years was inversely related to latitude. That is, plant communities in northern lakes were more persistent than communities in southern lakes. When we used multiple regression to find the…

Data from: A mechanistic underpinning for sigmoid dose-dependent infection

Theoretical models of environmentally transmitted diseases often assume that transmission is a constant process, which scales linearly with pathogen dose. Here we question the applicability of such an assumption and propose a sigmoidal form for the pathogens infectivity response. In our formulation, this response arises under two assumptions: 1) multiple invasion events are required for a successful pathogen infection and 2) the host invasion state is reversible. The first assumption reduces pathogen infection rates at low pathogen doses, while the second assumption, due to host immune function, leads to a saturating infection rate at high doses. The derived pathogen dose:infection rate -re…

Additional file 3: of Outside-host phage therapy as a biological control against environmental infectious diseases

Figure S2 Local stability analysis for susceptible host, infected host, pathogen and bacteriophage dynamics (S-I-P-F) when the phage transmission and release rate are varied (10− 9 0 is locally stable for high values of Λ F and β F (blue area). The coexistence equilibrium does not exist for lower values of when Λ F and β F (yellow area). In this case S and I become asymptotically extinct. Pathogen P always survives and even coexists with the phage F in the absence of the host S. The parameter values used are shown in Table 1. (TIF 2096 kb)

Additional file 4: of Outside-host phage therapy as a biological control against environmental infectious diseases

Figure S3 Time dynamics of pathogen, susceptible, infected host and bacteriophage dynamics (S-I-P-F) when the phage transmission and release rate are high (β F = 10− 3 and Λ F = 100). X-axis shows time in days and y-axis the population density. The pathogen population decreases close to extinction almost immediately, while there is a delay of approximately 47 days for the elimination of the disease and more than 50 days for decrease of the phage population close to extinction. The parameter values used are shown in Table 1. (TIF 2818 kb)

Additional file 1: of Outside-host phage therapy as a biological control against environmental infectious diseases

Figure S1 Time dynamics of pathogen, susceptible and infected host dynamics (S-I-P) in the absence of viral infection by bacteriophages and competition between the phage-resistant bacteria and the pathogen. X-axis and y-axis show time in days and the population density, respectively. Parameter values used are given in Table 1. In the absence of bacteriophages and phage-resistant bacteria, the pathogen (P), drives the susceptible host quickly extinct after which the infected host population goes extinct as well and the pathogen population stabilizes to grow saprothrophically in the absence of susceptible hosts. (TIF 2383 kb)