Energy storage and the evolution of population dynamics

We explore the mutual dependence of life history evolution and population dynamics by modeling a structured rotifer population that preys on a dynamic food supply. We focus on the ecological role of energy storage. A physiologically based submodel describes how individual predators allocate assimilated energy among growth, reproduction, and storage. We use invasibility analyses to predict evolutionary stable strategies for energy allocation. Various proxy measures of fitness based on measurable biological quantities, such as average population size or average per-capita fecundity, fail to predict evolutionary stable strategies. The predicted strategies indicate that selection strongly favors storage allocation for juveniles, but only for adults when prey densities are high. With the evolution of energy storage, population dynamics can shift from aperiodic to stable cycles without any need to invoke group selection.     

Approximation of a physiologically structured population model with seasonal reproduction by a stage-structured biomass model

Seasonal reproduction causes, due to the periodic inflow of young small individuals in the population, seasonal fluctuations in population size distributions. Seasonal reproduction furthermore implies that the energetic body condition of reproducing individuals varies over time. Through these mechanisms, seasonal reproduction likely affects population and community dynamics. While seasonal reproduction is often incorporated in population models using discrete time equations, these are not suitable for size-structured populations in which individuals grow continuously between reproductive events. Size-structured population models that consider seasonal reproduction, an explicit growing season and individual-level energetic processes exist in the form of physiologically structured population models. However, modeling large species ensembles with these models is virtually impossible. In this study, we therefore develop a simpler model framework by approximating a cohort-based size-structured population model with seasonal reproduction to a stage-structured biomass model of four ODEs. The model translates individual-level assumptions about food ingestion, bioenergetics, growth, investment in reproduction, storage of reproductive energy, and seasonal reproduction in stage-based processes at the population level. Numerical analysis of the two models shows similar values for the average biomass of juveniles, adults, and resource unless large-amplitude cycles with a single cohort dominating the population occur. The model framework can be extended by adding species or multiple juvenile and/or adult stages. This opens up possibilities to investigate population dynamics of interacting species while incorporating ontogenetic development and complex life histories in combination with seasonal reproduction.     

Interactions between year classes in the benthic amphipod Monoporeia affinis: effects on juvenile survival and growth

Summary The effect of 1-year-old Monoporeia (=Pontoporeia) affinis on the survival and growth of young of the year was studied in laboratory experiments. Amphipods were kept in jars with sediment and a continuous supply of cooled water for 3 months. Adults or juveniles were added to a control density of juveniles. Juvenile mortality increased at higher total densities, and was greater when adults were present. The mean length and dry weight of juveniles decreased at higher densities, but adults did not have a greater effect than juveniles. These density-dependent effects are probably caused by competition for food within and between year classes. Thus, a high abundance of juveniles or adults could reduce the recruitment of juveniles into the adult year-class the following year, and the suppression of juvenile growth could lower fecundity. The results of this experiment support the hypothesis, inferred from field data from the Baltic Sea and Swedish lakes, that intraspecific interactions contribute to fluctuations in the abundance of populations of M. affinis.     

Bifurcation theory, adaptive dynamics and dynamic energy budget-structured populations of iteroparous species

In this paper, we describe a technique to evaluate the evolutionary dynamics of the timing of spawning for iteroparous species. The life cycle of the species consists of three life stages, embryonic, juvenile and adult whereby the transitions of life stages (gametogenesis, birth and maturation) occur at species-specific sizes. The dynamics of the population is studied in a semi-chemostat environment where the inflowing food concentration is periodic (annual). A dynamic energy budget-based continuous-time model is used to describe the uptake of the food, storage in reserves and allocation of the energy to growth, maintenance, development (embryos, juveniles) and reproduction (adults). A discrete-event process is used for modelling reproduction. At a fixed spawning date of the year, the reproduction buffer is emptied and a new cohort is formed by eggs with a fixed size and energy content. The population consists of cohorts: for each year one consisting of individuals with the same age which die after their last reproduction event. The resulting mathematical model is a finite-dimensional set of ordinary differential equations with fixed 1-year periodic boundary conditions yielding a stroboscopic map. We will study the evolutionary development of the population using the adaptive dynamics approach. The trait is the timing of spawning. Pairwise and mutual invasibility plots are calculated using bifurcation analysis of the stroboscopic map. The evolutionary singular strategy value belonging to the evolutionary endpoint for the trait allows for an interpretation of the reproduction strategy of the population. In a case study, parameter values from the literature for the bivalve Macoma balthica are used.     

Niche overlap between herbivorous cladocerans; the role of food quality and habitat homogeneity

In spite of resource limitation, five abundant species of herbivorous metazoan zooplankton in a humic lake exhibited extensive niche overlap both with regard to seasonal and spatial occurrence, time of reproductive maxima, juvenile release and food choice. Their coexistence could not be explained by modifying predation, environmental oscillations or recolonization.Laboratory bottle experiments indicated only weak interspecific interactions between the three tested species at low food levels, but negative interactions were induced at elevated food levels. Bosmina appeared as competitively inferior during enrichment with cultured algae, but as the superior species during starvation. At low nutrient levels, all species coexisted for several generations with low reproduction, in accordance with the lake situation. It was concluded that the observed niche overlaps would be promoted if; 1) Intraspecific competition is more important than interspecific competition. 2) All species are co-adapted to low nutrient availability, food is quantitatively in surplus, but qualitatively deficient (mainly recycled detritus). During such conditions, no species would be capable of obtaining a population increase until extinction of the other species. This situation may be typical of oligotrophic humic lakes, and of other localities with low predation pressure and high inputs of allochthonous particulate carbon.     

Bridging Developmental Boundaries: Lifelong Dietary Patterns Modulate Life Histories in a Parthenogenetic Insect

Determining the effects of lifelong intake patterns on performance is challenging for many species, primarily because of methodological constraints. Here, we used a parthenogenetic insect (Carausius morosus) to determine the effects of limited and unlimited food availability across multiple life-history stages. Using a parthenogen allowed us to quantify intake by juvenile and adult females and to evaluate the morphological, physiological, and life-history responses to intake, all without the confounding influences of pair-housing, mating, and male behavior. In our study, growth rate prior to reproductive maturity was positively correlated with both adult and reproductive lifespans but negatively correlated with total lifespan. Food limitation had opposing effects on lifespan depending on when it was imposed, as it protracted development in juveniles but hastened death in adults. Food limitation also constrained reproduction regardless of when food was limited, although decreased fecundity was especially pronounced in individuals that were food-limited as late juveniles and adults. Additional carry-over effects of juvenile food limitation included smaller adult size and decreased body condition at the adult molt, but these effects were largely mitigated in insects that were switched to ad libitum feeding as late juveniles. Our data provide little support for the existence of a trade-off between longevity and fecundity, perhaps because these functions were fueled by different nutrient pools. However, insects that experienced a switch to the limited diet at reproductive maturity seem to have fueled egg production by drawing down body stores, thus providing some evidence for a life-history trade-off. Our results provide important insights into the effects of food limitation and indicate that performance is modulated by intake both within and across life-history stages.     

Year class splitting in the woodlouse Philoscia muscorum explained through studies of growth and survivorship

Year class splitting has been observed in a population of the terrestrial isopod Philoscia muscorum (Scopoli) in a fixed dune grassland. In this population, individuals born in one season differentiate into two forms, one of which grows to maturity in one year, the other in two.
Laboratory investigations of growth, fecundity and survivorship of individuals showed fundamental positive relationships between rate of growth and the ambient temperature/day light regime and between parental size and the number of offspring produced. Size also affected survivorship, with adults surviving better than juveniles. A link between survivorship and the temperature/day light regime was only obvious in small juveniles.
No direct physiological cause for the year class split could be derived from the laboratory studies, but a trade-off between the need for delayed or early breeding could result in the patterns of growth and reproduction observed in the field population. Delayed breeding allows continued growth of adults and hence larger numbers of offspring. Early breeding allows more growth of juveniles which greatly improves their chances of winter survival.     

Food limitation and body size in the life cycles of planktonic rotifers and cladocerans

This review considers what is known about the effects of food limitation upon the life cycle characteristics of rotifers and planktonic cladocerans. The characteristics considered in rotifers are the size of eggs, juveniles and adults and the durations of the juvenile phase and period of egg production. In cladocerans, the life history features dealt with are their length-weight relationships, the body size, instar stage, age and fecundity of the primiparous female and their fecundity-adult size relationship. The influence of limiting food conditions is demonstrated for these characteristics by comparison with the situation in non-limiting circumstances; the comparison is confined to experiments where food concentrations are quantified. A direct comparison is made between rotifers and cladocerans in conditions of defined food resource availability in terms of their length-weight relationships, the daily allocation of adults or near-adults to growth and reproduction and their threshold food concentrations. These comparisons are discussed in relation to the following topics: the high cost of cumulated respiration resulting from prolongation of the juvenile phase of body growth; the fundamentally different nature of growth in the two taxonomic groups; the body size of species and the size that must be attained for reproduction; the ecological implications of the very different threshold food concentrations.     

Intra-specific competition and density dependent juvenile growth

A difference equation model for the dynamics of a semelparous size-structured species consisting of juvenile and adult individuals is derived and studied. The adult population consists of two size classes, a smaller class and a larger more fertile class. Negative feedback occurs through slowed juvenile growth due to increased total population levels during the developmental period and consequently a smaller adult size at maturation. Intra-specific competition coefficients are size dependent and measure the strength of intra-specific competition between juveniles and adults. It is shown that equilibrium states in which adults and juveniles occur together at all times are in general destabilized by significantly increased juvenilevs adults competition with the result that stable periodic cycles appear, in which the generations alternate in time and hence avoid competition. This result supports the tenet that intra-specific competition between juveniles and adults is destabilizing. Exceptions to this destabilization principle are found, however, in which populations exhibiting non-equilibrium, aperiodic dynamics can be equilibrated by increase competition between juveniles and adults. This occurs, for example, when adult fertility and competition coefficients are significantly size class dependent. The author gratefully acknowledges the support of the Applied Mathematics Division and the Population Biology/Ecology Division of the National Science Foundation under NSF grant No. DMS-8902508. Research supported by the Department of Energy under contracts W-7405-ENG-36 and KC-07-01-01.     

Fluctuations in numbers in box populations of Drosophila and selective genetic mechanism of their regulation

Box populations of Drosophila melanogaster are characterized by two types of periodical fluctuations of numbers: with low and high frequency. High frequency fluctuations are determined by existence of preimago and imago stages and subsequent delay in density-dependent limitation of imago reproduction, duration of which is determined by time of preimago stage. The period of these fluctuations should be limited within two generation, that is confirmed by experimental data. Low frequency fluctuations with the period of 13-15 generations are the result of ecological density-dependent effect. In this case during pick density one can observe continuous degradation of population (i.e. decrease in fecundity and life time of imago) and following decrease in numbers. Temporary changes in fecundity of females and their offspring of the second generation are positively correlated with low frequency fluctuations in numbers. Such relationships show the possibility of density-dependent, cyclic, genetic changes in fecundity connected with fluctuations in numbers. It means that at the phase of growth in numbers when the density is still low, the selection is directed to the individuals with high fecundity sensible to overpopulation. The phase of decline in numbers is connected with high density and selection directed to the individuals with low fecundity in low density populations. The changes in genetic structure of fluctuating population lead to the weakening of this fluctuations and to the maintaining of population under such conditions.     

Experimental exposure of juvenile snails (Potamopyrgus antipodarum ) to infection by trematode larvae (Microphallus sp.): infectivity, fecundity compensation and growth

Host-parasite interactions that result in host castration are evolutionarily similar to predator-prey interactions because both interactions terminate reproduction for the host or prey. Yet, host-parasite interactions differ from predator-prey interactions in that infected hosts remain alive and potentially can make adjustments to their life-history strategy before castration is complete. Here we exposed juvenile snails (Potamopyrgus antipodarum) to infection by a digenetic trematode (Microphallus sp.) in order to determine whether: (1) pre-reproductive individuals could be infected, (2) individuals that were exposed to infection shifted resources to early reproduction (fecundity compensation), and (3) infected individuals exhibit altered growth rates relative to uninfected individuals. We found that juveniles are susceptible to infection; hence P. antipodarum could be selected for earlier maturation in populations where the risk of infection is high. We also found that fecundity compensation does not occur in this snail. Finally, we found that Microphallus-infected snails exhibit altered growth rates; individuals infected as juveniles have lower growth rates and are smaller than uninfected snails. These results suggest that growth is altered by infection of a trematode parasite but reproduction in uninfected snails is not induced by exposure to trematode eggs. Received: 11 January 1998 / Accepted: 19 May 1998     

Reproductive effort in viscous populations

Abstract.— Here I study a kin selection model of reproductive effort, the allocation of resources to fecundity versus survival, in a patch-structured population. Breeding females remain in the same patch for life. Offspring have costly, partial long-distance dispersal and compete for breeding sites, which become vacant upon the death of previous occupants. The main result is that the evolutionarily stable reproductive effort decreases as offspring dispersal rate increases. The result can be understood as follows: In a well-mixed population with global competition, neither adults nor juveniles compete with relatives, but in a patch-structured population with dispersal restricted to the juvenile phase, juveniles experience relatively less competition with relatives than adults, thus making juveniles relatively more valuable. Because this asymmetry between adults and juveniles decreases with the dispersal rate, so does the evolutionarily stable level of allocation to fecundity.     

Alternative dynamical states in stage-structured consumer populations

The population dynamics of a consumer population with an internal structure is investigated. The population is divided into juvenile and adult individuals that consume different resources and do not interfere with each other. Over a broad range of external conditions (varying mortality and different resource levels), alternative stable states exist. These population states correspond to domination of juveniles and domination of adults, respectively. When mortality is varied, hysteresis between the alternative states only occurs if juveniles have more resources than adults. In the opposite case the juvenile-dominated state is stable for all values of mortality, but the adult-dominated state is not. When the population is modelled with more than one juvenile stage, the adult-dominated state becomes a periodic orbit due to a delay in the regulatory mechanism of the population dynamics. It is shown numerically that the stage-structured model converges to a model with continuous size structure for very large numbers of successive juvenile stages.     

Simplifying a physiologically structured population model to a stage-structured biomass model

We formulate and analyze an archetypal consumer-resource model in terms of ordinary differential equations that consistently translates individual life history processes, in particular food-dependent growth in body size and stage-specific differences between juveniles and adults in resource use and mortality, to the population level. This stage-structured model is derived as an approximation to a physiologically structured population model, which accounts for a complete size-distribution of the consumer population and which is based on assumptions about the energy budget and size-dependent life history of individual consumers. The approximation ensures that under equilibrium conditions predictions of both models are completely identical. In addition we find that under non-equilibrium conditions the stage-structured model gives rise to dynamics that closely approximate the dynamics exhibited by the size-structured model, as long as adult consumers are superior foragers than juveniles with a higher mass-specific ingestion rate. When the mass-specific intake rate of juvenile consumers is higher, the size-structured model exhibits single-generation cycles, in which a single cohort of consumers dominates population dynamics throughout its life time and the population composition varies over time between a dominance by juveniles and adults, respectively. The stage-structured model does not capture these dynamics because it incorporates a distributed time delay between the birth and maturation of an individual organism in contrast to the size-structured model, in which maturation is a discrete event in individual life history. We investigate model dynamics with both semi-chemostat and logistic resource growth.     

Ontogenetic changes in Mammalian feeding: insights from electromyographic data

All infant mammals make a transition from suckling milk to eating solid foods. Yet, the neuromuscular implications of the transition from a liquid-only diet to solid foods are unknown even though the transport and swallowing of liquids is different from that of solids. We used legacy electromyography (EMG) data to test hypotheses concerning the changes in motor pattern and neuromuscular control that occur during the transition from an all-liquid diet to consumption of solid food in a porcine model. EMG signals were recorded from five oropharyngeal muscles in pigs at three developmental stages (infants, juveniles, and adults) feeding on milk, on food of an intermediate consistency (porridge), and on dry chow (juveniles and adults only). We measured cycle frequency and its variation in "transport cycles" and "swallow cycles". In the swallow cycles, a measure of variation of the EMG signal was also calculated. Variation in cycle frequency for transport and swallow cycles was lowest in adults, as predicted, suggesting that maturation of feeding mechanisms occurs as animals reach adulthood. Infants had lower variation in transport cycle frequency than did juveniles drinking milk, which may be due to the greater efficiency of the infant's tight oral seal against the teat during suckling, compared to a juvenile drinking from a bowl where a tight seal is not possible. Within juveniles, variation in both transport and swallow cycle frequencies was directly related to food consistency, with the highest variation occurring when drinking milk and the lowest when feeding on solid food. There was no difference in the variation of the EMG activity between intact infants and juveniles swallowing milk, although when the latter swallow porridge the EMG signals were less variable than for milk. These results suggest that consistency of food is a highly significant determinant of the variation in motor pattern, particularly in newly weaned animals.     

Absence of predation eliminates coexistence: experience from the fish–zooplankton interface

Examples from fishless aquatic habitats show that competition among zooplankton for resources instigates rapid exclusion of competitively inferior species in the absence of fish predation, and leads to resource monopolization by the superior competitor. This may be a single species or a few clones with large body size: a cladoceran such as Daphnia pulicaria, or a branchiopod such as Artemia franciscana, each building its population to a density far higher than those found in habitats with fish. The example of zooplankton from two different fish-free habitats demonstrates the overpowering force of fish predation by highlighting the consequences of its absence. Released from the mortality caused by predation, a population of a superior competitor remains at a density equal to the carrying capacity of its habitat, in a steady state with its food resources, consisting of small green flagellate algae, which are successful in compensating high loss rates due to grazing, by fast growth. In such a situation, the high filtering rate of Daphnia or Artemia reduces resources to levels that are sufficient for assimilation to cover the costs of respiration (threshold food concentration) in adults but not in juveniles. This implies long periods of persistence of adults refraining from producing live young, because production of instantly hatching eggs would be maladaptive. Severe competition for limiting resources imposes a strong selective pressure for postponing reproduction or for producing resting eggs until food levels have increased. Offspring can only survive when born in a short time window between such an increase in food levels and its subsequent decline resulting from population growth and intense grazing by juveniles. Such zooplanktons become not only a single-species community, but also form a single cohort with a long-lifespan population. The observations support the notion that diversity may be sustained only where predation keeps densities of coexisting species at levels much below the carrying capacity, as suggested by Hutchinson 50 years ago.     

Experimental evidence of size-selective harvest and environmental stochasticity effects on population demography,fluctuations and non-linearity

Theory and analyses of fisheries data sets indicate that harvesting can alter population structure and destabilise non-linear processes, which increases population fluctuations. We conducted a factorial experiment on the population dynamics of Daphnia magna in relation to size-selective harvesting and stochasticity of food supply. Harvesting and stochasticity treatments both increased population fluctuations. Timeseries analysis indicated that fluctuations in control populations were non-linear, and non-linearity increased substantially in response to harvesting. Both harvesting and stochasticity induced population juvenescence, but harvesting did so via the depletion of adults, whereas stochasticity increased the abundance of juveniles. A fitted fisheries model indicated that harvesting shifted populations towards higher reproductive rates and larger-magnitude damped oscillations that amplify demographic noise. These findings provide experimental evidence that harvesting increases the non-linearity of population fluctuations and that both harvesting and stochasticity increase population variability and juvenescence.     

Body size and food thresholds for zero growth in Dreissena polymorpha: a mechanism underlying intraspecific competition

1. Dreissena polymorpha is an extraordinarily successful invasive species that shows high recruitment of small juvenile mussels on established mussel banks. Such juvenile settlement on, and overgrowth of, large adult mussels; however, leads to competition with adults, and often at high densities and low‐food concentrations. 2. The concept of food thresholds for zero growth has been a powerful approach to explaining size‐related exploitative competition in different zooplankton species. We applied it to investigate whether food threshold concentrations for zero growth (C₀) differ between juvenile and adult zebra mussels. 3. By determining body mass growth at various concentrations of a diet mixture (Nannochloropsis limnetica and Isochrysis aff. galbana) we demonstrate that the threshold food concentration for growth of juvenile mussels (C₀ = 0.08 mg C L⁻¹) is substantially lower than that for adults (C₀ = 0.36 mg C L⁻¹). 4. This indicates that, at low food availability, juvenile zebra mussels are competitively superior to their larger conspecifics. Within zebra mussel banks plankton food is substantially depleted and so the observed mechanism might ensure juvenile success and therefore the regeneration of mussel banks in nature.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Simultaneous effects of food limitation and inducible resistance on herbivore population dynamics

Many herbivore populations fluctuate temporally, but the causes of those fluctuations remain unclear. Plant inducible resistance can theoretically cause herbivore population fluctuations, because herbivory may induce plant changes that reduce the survival or reproduction of later-feeding herbivores. Herbivory can also simply reduce the quantity of food available for later feeders and this, too, can cause population fluctuations. Inducible resistance and food limitation often occur simultaneously, yet whether they jointly facilitate or suppress herbivore fluctuations remains largely unexplored. We present models that suggest that food limitation and inducible resistance may have synergistic effects on herbivore population dynamics. The population-level response of the food plant to herbivory and the details of how inducible resistance affects herbivore performance both influence the resulting herbivore dynamics. Our results identify some biological properties of plant-herbivore systems that might determine whether or not cycles occur, and suggest that future empirical and theoretical population dynamics studies should account for the effects of both food limitation and inducible resistance.