Disentangling demographic co‐effects of predation and pollution on population dynamics

In nature species react to a variety of endogenous and exogenous ecological factors. Understanding the mechanisms by which these factors interact and drive population dynamics is a need for understanding and managing ecosystems. In this study we assess, using laboratory experiments, the effects that the combinations of two exogenous factors exert on the endogenous structure of the population dynamics of a size‐structured population of Daphnia. One exogenous factor was size‐selective predation, which was applied on experimental populations through simulating: 1) selective predation on small prey, 2) selective predation on large prey and 3) non‐selective predation. The second exogenous factor was pesticide exposure, applied experimentally in a quasi‐continuous regime. Our analysis combined theoretical models and statistical testing of experimental data for analyzing how the density dependence structure of the population dynamics was shifted by the different exogenous factors. Our results showed that pesticide exposure interacted with the mode of predation in determining the endogenous dynamics. Populations exposed to the pesticide and to either selective predation on newborns or selective predation on adults exhibited marked nonlinear effects of pesticide exposure. However, the specific mechanisms behind such nonlinear effects were dependent on the mode of size‐selectivity. In populations under non‐selective predation the pesticide exposure exerted a weak lateral effect. The ways in which endogenous process and exogenous factors may interact determine population dynamics. Increases in equilibrium density results in higher variance of population fluctuations but do not modify the stability properties of the system, while changes in the maximum growth rate induce changes in the dynamic regimes and stability properties of the population. Future consideration for research includes the consequences of the seasonal variation in the composition and activity of the predator assembly in interaction with the seasonal variation in exposure to agrochemicals on freshwater population dynamics.     

Climate effects and feedback structure determining weed population dynamics in a long-term experiment

Pest control is one of the areas in which population dynamic theory has been successfully applied to solve practical problems. However, the links between population dynamic theory and model construction have been less emphasized in the management and control of weed populations. Most management models of weed population dynamics have emphasized the role of the endogenous process, but the role of exogenous variables such as climate have been ignored in the study of weed populations and their management. Here, we use long-term data (22 years) on two annual weed species from a locality in Central Spain to determine the importance of endogenous and exogenous processes (local and large-scale climate factors). Our modeling study determined two different feedback structures and climate effects in the two weed species analyzed. While Descurainia sophia exhibited a second-order feedback and low climate influence, Veronica hederifolia was characterized by a first-order feedback structure and important effects from temperature and rainfall. Our results strongly suggest the importance of theoretical population dynamics in understanding plant population systems. Moreover, the use of this approach, discerning between the effect of exogenous and endogenous factors, can be fundamental to applying weed management practices in agricultural systems and to controlling invasive weedy species. This is a radical change from most approaches currently used to guide weed and invasive weedy species managements.     

Evolution of resistance to anti-cancer therapy during general dosing schedules

Anti-cancer drugs targeted to specific oncogenic pathways have shown promising therapeutic results in the past few years; however, drug resistance remains an important obstacle for these therapies. Resistance to these drugs can emerge due to a variety of reasons including genetic or epigenetic changes which alter the binding site of the drug target, cellular metabolism or export mechanisms. Obtaining a better understanding of the evolution of resistant populations during therapy may enable the design of more effective therapeutic regimens which prevent or delay progression of disease due to resistance. In this paper, we use stochastic mathematical models to study the evolutionary dynamics of resistance under time-varying dosing schedules and pharmacokinetic effects. The populations of sensitive and resistant cells are modeled as multi-type non-homogeneous birth-death processes in which the drug concentration affects the birth and death rates of both the sensitive and resistant cell populations in continuous time. This flexible model allows us to consider the effects of generalized treatment strategies as well as detailed pharmacokinetic phenomena such as drug elimination and accumulation over multiple doses. We develop estimates for the probability of developing resistance and moments of the size of the resistant cell population. With these estimates, we optimize treatment schedules over a subspace of tolerated schedules to minimize the risk of disease progression due to resistance as well as locate ideal schedules for controlling the population size of resistant clones in situations where resistance is inevitable. Our methodology can be used to describe dynamics of resistance arising due to a single (epi)genetic alteration in any tumor type.     

Climate mediated exogenous forcing and synchrony in populations of the oak aphid in the UK

Contemporary population dynamics theory suggests that animal fluctuations in nature are the result of the combined forces of intrinsic and exogenous factors. Weather is the iconic example of an exogenous force. The common approach for analyzing the relationship between population size and climatic variables is by simple correlation or using the climate as an additive covariable in statistical models. Here, we evaluated different functional forms in which climatic variables could influence population dynamics of the oak aphid Tuberculatus annulatus both in each locality and in relation to synchrony between localities. Results indicate that in at least four of eight aphid populations, climate influences population dynamics by modifying the carrying capacity of the system (lateral effect mediated by winter precipitation). Additionally, path analysis showed that synchrony in population dynamics is highly correlated with synchrony in winter precipitation regime, and the spatial scale of both processes is similar, which suggests that this is an example of the Moran effect. Our results show the key effects of precipitation on intra and inter population processes of this aphid. The methods used, mixing population dynamics modelling and test of synchrony, allowed us to connect the direct and indirect effects of exogenous variables into each population with patterns of synchrony inter populations.     

Resonance effects and outbreaks in ecological time series

Organismal response to environmental variability is an important aspect of ecological processes. We propose new mechanisms whereby environmental variability can cause cyclic population outbreaks due to the nonlinearity of the organismal response. We consider stage-structured populations that respond to variable environments with variable diapause or dormancy, and in which cyclic changes of the environment induce a resonance-like boost in the population size. If there is also a stochastic component of variation in the environment, the population outbreaks are magnified by the phenomenon of "stochastic resonance". The results show that large population fluctuations may not be due to extrinsic or intrinsic factors alone, but to a nonlinear interaction between the external environment and internal population processes. Indeed, in the presence of such nonlinearities even very small environmental fluctuations can cause massive fluctuations in population size. Our theoretical results may help to explain periodic population cycles and outbreak dynamics found in many infectious diseases and pest species. We also discuss the evolution of the response parameters that regulate diapause or dormancy and promote the outbreak dynamics in variable environments.     

Population dynamics of two rodent species in agro-ecosystems of central Argentina: intra-specific competition,land-use,and climate effects

Understanding the role of feedback structure (endogenous processes) and exogenous (climatic and environmental) factors in shaping the dynamics of natural populations is a central challenge within the field of population ecology. We attempted to explain the numerical fluctuations of two sympatric rodent species in agro-ecosystems of central Argentina using Royama’s theoretical framework for analyzing the dynamics of populations influenced by exogenous climatic forces. We found that both rodent species show a first-order negative feedback structure, suggesting that these populations are regulated by intra-specific competition (limited by food, space, or enemy-free space). In Akodon azarae endogenous structure seems to be very strongly influenced by human land-use represented by annual minimum normalized difference vegetation index (NDVI), with spring and summer rainfall having little influence upon carrying capacity. Calomys venustus’ population dynamics, on the other hand, seem to be more affected by local climate, also with spring and summer rainfall influencing the carrying capacity of the environment, but combined with spring mean temperature. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Non-linear feedback processes and a latitudinal gradient in the climatic effects determine green spruce aphid outbreaks in the UK

The role of climatic fluctuations in determining the dynamics of insect populations has been a classical problem in population ecology. Here, we use long-term annual data on green spruce aphid populations at nine localities in the UK for determining the importance of endogenous processes, local weather and large-scale climatic factors. We rely on diagnostic and modelling tools from population dynamic theory to analyse these long-term data and to determine the role of the North Atlantic Oscillation (NAO) and local weather as exogenous factors influencing aphid dynamics. Our modelling suggests that the key elements determining population fluctuations in green spruce aphid populations in the UK are the strong non-linear feedback structure, the high potential for population growth and the effects of winter and spring weather. The results indicate that the main effect of the NAO on green spruce aphid populations is operating through the effect of winter temperatures on the maximum per capita growth rate (R_m). In particular, we can predict quite accurately the occurrence of an outbreak by using a simple logistic model with weather as a perturbation effect. However, model predictions using different climatic variables showed a clear geographical signature. The NAO and winter temperature were best for predicting observed dynamics toward the southern localities, while spring temperature was a much better predictor of aphid dynamics at northern localities. Although aphid species are characterized by complex life-cycles, we emphasize the value of simple and general population dynamic models in predicting their dynamics.     

The relative roles of density and climatic variation on population dynamics and fecundity rates in three contrasting ungulate species

The relative influences of density-dependent and -independent processes on vital rates and population dynamics have been debated in ecology for over half a century, yet it is only recently that both processes have been shown to operate within the same population. However, generalizations on the role of each process across species are rare. Using a process-orientated generalized linear modelling approach we show that variations in fecundity rates in populations of three species of ungulates with contrasting life histories are associated with density and winter weather in a remarkably similar manner. However, there are differences and we speculate that they are a result of differences in size between the species. Much previous research exploring the association between vital rates, population dynamics and density-dependent and -independent processes has used pattern-orientated approaches to decompose time-series into contributions from density-dependent and -independent processes. Results from these analyses are sometimes used to infer associations between vital rates, density and climatic variables. We compare results from pattern-orientated analyses of time-series with process-orientated analyses and report that the two approaches give different results. The approach of analysing relationships between vital rates, density and climatic variables may detect important processes influencing population dynamics that time-series methodologies may overlook.     

Population cycles produced by delayed density dependence in an annual plant

In contrast to insect and animal populations, little attention has been directed to the study of cycles in plant populations. It has been argued on theoretical grounds that plants present stable dynamics. Nevertheless, there are examples where plant populations appear to exhibit oscillatory dynamics, but the oscillatory signal is variable and comes from very short time series data. Using a combination of time series, models, and empirical results, we present evidence of population cycles for Descurania sophia in a 16-year field experiment. Endogenous and exogenous causal mechanisms were studied to identify processes underlying this temporal dynamic. Our results show a 4-year cycle produced by delayed density dependence. We suggest that high nutrient levels might be responsible for the observed dynamics of D. sophia. Our results suggest that although plant population dynamics may be stabilized by direct density dependence, delayed density dependence could destabilize dynamics.     

Host-specificity of AM fungal population growth rates can generate feedback on plant growth

While the mutualistic interaction between plants and AM fungi is of obvious importance to ecosystem processes, the factors influencing the ecological and evolutionary dynamics within this interaction are poorly understood. The mutual interdependence of plant and AM fungal relative growth rates could generate complex dynamics in which the composition of the AM fungal community changes due to association with host and this change in fungal composition then differentially feeds back on plant growth. I first review evidence for feedback dynamics and then present an approach to evaluating such complex dynamics. I specifically present evidence of host-specific differences in the population growth rates of AM fungi. Pure cultures of AM fungi were mixed to produce the initial fungal community. This community was then distributed into replicate pots and grown with one of four co-occurring plant species. Distinct compositions of AM fungal spores were produced on different host species. The AM fungal communities were then inoculated back onto their own host species and grown for a second growing season. The differentiation observed in the first generation was enhanced during this second generation, verifying that the measure of spore composition reflects host-specific differences in AM fungal population growth rates. In further work on this system, I have found evidence of negative feedback through two pairs of plant species. The dynamic within the AM fungal community can thereby contribute to the coexistence of plant species.     

Population dynamics of coral-reef fishes: Controversial concepts and hypotheses

Abstract Knowledge of processes that drive the local population dynamics of coral-reef fishes is important for managing reef fisheries, and for using these species as models for understanding the ecology of demersal marine fishes in general. However, the reef-fish literature is replete with poorly defined concepts and vague hypotheses regarding the issue of population dynamics. Dichotomous arguments, such as whether or not recruitment drives population dynamics, are misdirected because they fail to incorporate several important concepts. First, changes in local population size are driven by four demographic rates (birth, death, immigration and emigration), all of which must be studied to understand population dynamics. Second, all populations that persist do so because at least one of these demographic rates operates in a density-dependent way that is both sufficiently strong and appropriately time-lagged. Therefore, identifying the source(s) of direct density dependence is critical for understanding the limits to variation in population size (i.e. population regulation). Third, regulation does not imply a simple point equilibrium in population size; density dependence in populations of reef fishes is bound to lie within a field of stochastic variation, and thus be difficult to detect. Since its formal origin in 1981, the ‘recruitment limitation’ hypothesis for explaining local population dynamics in reef fishes has undergone ambiguous changes in definition that threaten its usefulness. ‘Recruitment, ‘originally defined as the appearance of newly settled fish on a reef, more recently is often measured months after settlement, thus confounding pre- and post-settlement processes. ‘Limitation, ‘ which originally referred to recruitment being so low as to preclude local populations from reaching densities where resources were limiting, is more recently defined as an absence of any form of density dependence after settlement. The most effective means of testing whether post-settlement mortality is in fact density-independent is to examine patterns of mortality directly, rather than indirectly by interpreting the shape of the relationship between initial recruit density and subsequent adult density within a cohort (the recruit-adult function). Understanding the population dynamics of coral-reef fishes will require a more equitable focus on all four demographic rates, be they density dependent or not, as well as greater attention to identifying sources of density dependence. Such a pluralistic focus necessitates integrated studies of both pre- and post-settlement processes conducted at multiple spatial and temporal scales. For example, recent evidence suggests that density-dependent pre-dation on new recruits that have settled among reefs at different densities may prove to be an important source of local population regulation, especially via the aggregative response of transient piscivores.     

Individual variability and population regulation: a model of the significance of within-generation density dependence

Most models of theoretical population ecology consider population density as a state variable and thus ignore the fact that populations are composed not of identical average individuals but of individuals which are usually different. However, this individual variability may be important for population regulation. We therefore analysed an individual-based population model which explicitly describes within-generation processes, i.e. individual growth, starvation, and resource dynamics. The results show that if population dynamics are dominated by slow changes in resource level, the population size in the model undergoes wide oscillation, often leading to extinction. If, on the other hand, fast within-generation processes predominate, such as starvation and sudden drops in resource levels, the population fluctuates to a limited extent around an average. Within-generation density dependence may thus be an important mechanism which is largely ignored in classic time-discrete state-variable models. We conclude that the individual-based approach provides important insights into the hierarchical organization of population dynamics, i.e. the relationship between fast processes at the individual level and slower processes at the population level.     

Dynamic macroecology on ecological time-scales

Aim The discipline of macroecology is increasingly being regarded as an effective vehicle for the evaluation of recent population‐ to ecosystem‐level responses to widespread human and environmental influences. However, due to the prevalent use of time‐averaged and cumulative data in macroecological analyses, the majority of the patterns that emerge from research in this field can be regarded as static. Here we review the application of dynamic macroecological analyses to changes in relationships between macroecological variables on seasonal to decadal scales. We illustrate the strength of this perspective for documenting changing patterns and testing hypotheses related to these dynamics on ecological time‐scales. Location Studies were compiled and reviewed from terrestrial and aquatic ecosystems. Methods We review examples of temporal changes in macroecological patterns driven by recent anthropogenic influences and environmental change. Results The dynamic nature of macroecological patterns on ecological time‐scales has been revealed in recent years across a wide range of ecosystems, largely through the development, maintenance and analysis of biotic and environmental monitoring time series. The resultant analyses complement examinations of dynamics over evolutionary time and have similarly revealed that static portrayals can conceal important temporal dynamics that underlie the patterns of interest. As a consequence, static depictions, resting as they do on comparative analyses in which the validity of space‐for‐time substitutions is assumed, may be of limited use for testing hypotheses related to the mechanisms underlying the patterns revealed and, by extension, the development of reliable predictions of future states. Main conclusions Recent dynamic macroecological analyses have demonstrated the utility of combined spatial and temporal replication, and have contributed to hypothesis testing related to the mechanistic processes underlying changes in macroecological patterns on ecological time‐scales. We suggest four specific avenues of future research to further the development and application of temporal approaches on similar time‐scales within the field of macroecology.     

Genetic drift in an infinite population. The pseudohitchhiking model

Selected substitutions at one locus can induce stochastic dynamics that resemble genetic drift at a closely linked neutral locus. The pseudohitchhiking model is a one-locus model that approximates these effects and can be used to describe the major consequences of linked selection. As the changes in neutral allele frequencies when hitchhiking are rapid, diffusion theory is not appropriate for studying neutral dynamics. A stationary distribution and some results on substitution processes are presented that use the theory of continuous-time Markov processes with discontinuous sample paths. The coalescent of the pseudohitchhiking model is shown to have a random number of branches at each node, which leads to a frequency spectrum that is different from that of the equilibrium neutral model. If genetic draft, the name given to these induced stochastic effects, is a more important stochastic force than genetic drift, then a number of paradoxes that have plagued population genetics disappear.     

Temporal dynamics of a local fish community are strongly affected by immigration from the surrounding metacommunity

A 5‐year time series of annual censuses was collected from a large floodplain lake to determine how dynamics of the local fish community were affected by changes in hydrological connectivity with the surrounding metacommunity. The lake was disconnected from the metacommunity for 1 year prior to our study and remained disconnected until 3 months before our third annual census, when a flood reconnected the lake to the metacommunity. We determined how changes in connectivity affected temporal dynamics of (1) local community composition and (2) the population composition, condition, and growth of catfish, to shed light on how immigration of other species might affect local population dynamics. Before reconnection, the community was likely shaped by interactions between the local environment and species traits. The reconnection caused significant immigration and change in community composition and correlated with a significant and abrupt decline in catfish condition, growth, and abundance; effects likely due to the immigration of a competitor with a similar trophic niche: carp. The community was slow to return to its preconnection state, which may be due to dispersal traits of the fishes, and a time‐lag in the recovery of the local catfish population following transient intensification of species interactions. The dynamics observed were concordant with the species sorting and mass‐effects perspectives of metacommunity theory. Floods cause episodic dispersal in floodplain fish metacommunities, and so, flood frequency determines the relative importance of regional and local processes. Local processes may be particularly important to certain species, but these species may need sufficient time between floods for population increase, before the next flood‐induced dispersal episode brings competitors and predators that might cause population decline. Accordingly, species coexistence in these metacommunities may be facilitated by spatiotemporal storage effects, which may in turn be regulated by flood frequency.     

Weather variability impacts on oviposition dynamics of the southern house mosquito at intermediate time scales

Oviposition is a major event in the life history of mosquitoes, shaping both individual fitness and vectorial capacity. Several exogenous factors have been shown as important for the dynamic forcing of oviposition at finely (hourly) and coarsely (monthly or season to season) grained temporal scales. However, field studies addressing the interplay of weather factors on oviposition dynamics at the intermediate (days to weeks) time scale are missing. Here, we present the results from a field study that showed the oviposition dynamics of the southern house mosquito, Culex quinquefasciatus Say (Diptera: Culicidae), to be: (i) primarily dictated by relative humidity; and (ii) disrupted by rainfall events that resulted in a modified sensitivity to relative humidity. Rainfall changed the concentration of ammonia, a major limiting resource of microbes used as food by mosquito larvae. Following major rainfall events, the importance of relative humidity in forcing the oviposition dynamics also changed. Finally, our results indicate that qualitative changes in oviposition habitats modify the importance of weather variables as predictors of mosquito oviposition dynamics.     

Modelling memory functions with recurrent neural networks consisting of input compensation units: I. Static situations

Humans are able to form internal representations of the information they process—a capability which enables them to perform many different memory tasks. Therefore, the neural system has to learn somehow to represent aspects of the environmental situation; this process is assumed to be based on synaptic changes. The situations to be represented are various as for example different types of static patterns but also dynamic scenes. How are neural networks consisting of mutually connected neurons capable of performing such tasks? Here we propose a new neuronal structure for artificial neurons. This structure allows one to disentangle the dynamics of the recurrent connectivity from the dynamics induced by synaptic changes due to the learning processes. The error signal is computed locally within the individual neuron. Thus, online learning is possible without any additional structures. Recurrent neural networks equipped with these computational units cope with different memory tasks. Examples illustrate how information is extracted from environmental situations comprising fixed patterns to produce sustained activity and to deal with simple algebraic relations.     

On the changing concept of evolutionary population stability as a reflection of a changing point of view in the quantitative theory of evolution

 Eighteen different terms, currently employed to define various concepts of evolutionary stability in population dynamics are mentioned in this paper. Most of these terms are used in different connotations and even different meanings by different authors. On the other hand, different terms are often employed by different authors to define quite the same concept. Twenty-five years ago there was only one, well-defined, concept of stability, universally recognized in the field. In this paper I will try to relate the recent confusion, concerning concepts of population stability, with a more serious, though not that well-recognized, confusion in the modern analytic approach to population dynamics and quantitative evolution. Concepts of population stability will be examined in relation to each other on the one hand and, on the other hand, in relation to two dichotomies regarding the dynamic processes to which they correspond: Short-term versus long-term processes and processes concerning phenotypic changes versus process concerning genotypic changes. A hopefully more consistent use of the current terminology is suggested. Received 15 August 1993; received in revised form 15 September 1994     

Regional dynamics of a patchily distributed herbivore along an altitudinal gradient

Abstract.  1. Metapopulation dynamics should be more important at the borders of species distributions due to two main factors: (1) populations are less abundant and fluctuate more at the borders than in the centre of their distributions, and (2) resources in the range margins of species distributions are often more scarce and fragmented.
2. Most metapopulation studies have been performed in a fraction of the entire distribution of species. The main goal here is to study the population dynamics of a narrowly distributed species including both the borders and the centre of the distribution, and to test the predictions described above.
3. The density and extinction events in a patchily distributed species, Timarcha lugens , was quantified for 5 years along an altitudinal gradient including the upper and lower limits of the species distribution. The dispersal ability of Timarcha was also studied using a mark–release–recapture study.
4. Extinction events and empty patches were only found at the borders of Timarcha distribution. The fluctuation in beetle density was greater in patches suffering extinction events. Resource abundance was negatively related to beetle density and positively related to extinction events. In addition, the dispersal rate among patches was very low and beetles moved distances of no further than 25 m.
5. Population density governs the extinction events in this system, and its fluctuation was more evident near the border of the distribution. Both factors together with the relative population stability in patches at medium and high altitudes, and the low dispersal rate of the individuals support the idea of a source–sink metapopulation structure in T. lugens .