The impact of diapause on the evolution of other life history traits in flesh flies

Summary Flesh flies (Sarcophagidae) collected in Costa Rica and Panama lack the pupal diapause that is characteristic of flesh flies from the temperate zone and tropical Africa. The absence of a diapause capacity in the neotropical species correlates with several other life history traits: in most species the post feeding wandering phase of the third larval instar is longer and duration is more variable, adult life is longer, clutch size is smaller, and more clutches are produced. Among species that have the capacity for diapause, risk is invested primarily in the diapausing stage and other life stages are brief. Though diapausing species are short-lived, they produce as many or more progeny than nondiapausing species by increasing clutch size. The slower and more variable developmental rate and increased adult longevity desynchronizes development and permits the nondiapausing species to spread an environmental risk over different stages of the life cycle, thus offering an alternative to diapause. Other traits such as body size, developmental velocity, thermal constant thresholds, thermal constants, age at first reproduction, and the interval between clutches do not appear related to the capacity for diapause.     

The case for negative senescence

Negative senescence is characterized by a decline in mortality with age after reproductive maturity, generally accompanied by an increase in fecundity. Hamilton (1966) ruled out negative senescence: we adumbrate the deficiencies of his model. We review empirical studies of various plants and some kinds of animals that may experience negative senescence and conclude that negative senescence may be widespread, especially in indeterminate-growth species for which size and fertility increase with age. We develop optimization models of life-history strategies that demonstrate that negative senescence is theoretically possible. More generally, our models contribute to understanding of the evolutionary and demographic forces that mold the age-trajectories of mortality, fertility and growth.     

How can we model selectively neutral density dependence in evolutionary games

The problem of density dependence appears in all approaches to the modelling of population dynamics. It is pertinent to classic models (i.e., Lotka-Volterra's), and also population genetics and game theoretical models related to the replicator dynamics. There is no density dependence in the classic formulation of replicator dynamics, which means that population size may grow to infinity. Therefore the question arises: How is unlimited population growth suppressed in frequency-dependent models? Two categories of solutions can be found in the literature. In the first, replicator dynamics is independent of background fitness. In the second type of solution, a multiplicative suppression coefficient is used, as in a logistic equation. Both approaches have disadvantages. The first one is incompatible with the methods of life history theory and basic probabilistic intuitions. The logistic type of suppression of per capita growth rate stops trajectories of selection when population size reaches the maximal value (carrying capacity); hence this method does not satisfy selective neutrality. To overcome these difficulties, we must explicitly consider turn-over of individuals dependent on mortality rate. This new approach leads to two interesting predictions. First, the equilibrium value of population size is lower than carrying capacity and depends on the mortality rate. Second, although the phase portrait of selection trajectories is the same as in density-independent replicator dynamics, pace of selection slows down when population size approaches equilibrium, and then remains constant and dependent on the rate of turn-over of individuals.     

The effects of background mortality on optimal reproduction in a seasonal environment

We consider optimal annual routines of reproductive behaviour in a seasonal environment. In our model the condition of the organism is adversely affected by hard work, but can recover during easy periods. Our analysis concentrates on the effects of background mortality (i.e., mortality that cannot be avoided) on the optimal strategy and how often an organism following this strategy breeds. In particular, we are concerned with whether reproduction occurs at specific times of year (entrained to the annual cycle), and if so then how many reproductive bouts occur per year. We find that an increase in background mortality can have various effects. If the animal is entrained to the annual cycle and has one breeding attempt per year, then breeding tends to occur earlier and there may be two breeding attempts per season. Another possible outcome is that breeding is no longer entrained. If the animal is entrained but sometimes skips reproduction so that it does not breed every year, then an increase in mortality may make it more likely that the animal breeds every year. We show that as background mortality increases the resultant increase in the frequency of breeding contributes to the increase in annual mortality. We also explore the effects of mortality on the timing of reproduction within a year, highlighting the tension between the interests of the parent and that of the young.     

The co-evolution of intergenerational transfers and longevity: an optimal life history approach

How would resources be allocated among fertility, survival, and growth in an optimal life history? The budget constraint assumed by past treatments limits the energy used by each individual at each instant to what it produces at that instant. We consider under what conditions energy transfers from adults, which relax the rigid constraint by permitting energetic dependency and faster growth for the offspring, would be advantageous. In a sense, such transfers permit borrowing and lending across the life history. Higher survival and greater efficiency in energy production at older ages than younger both favor the evolution of transfers. We show that if such transfers are advantageous, then increased survival up to the age of making the transfers must co-evolve with the transfers themselves.     

The spatial characteristics of stand structure in Pinus torreyana

The arrangement of trees within a stand by location and age (stand structure) is in part determined by the life history strategy of the species and the disturbance history of the stand. In western North America such disturbances are often the product of wildfires and human management activities. The current study uses spatial analysis to characterize three stands of Pinus torreyana with known disturbance histories. Two stands are located at Torrey Pines State Reserve (TPSR). Of these, one stand has burned twice since 1972. Fire has been successfully excluded from the other stand since the early part of this century. A third stand, on Santa Rosa Island, Channel Islands National Park (SRI), has been grazed heavily since the mid-19th century and has not experienced fire since that time. One-hectare study plots were established in the interior of each stand. Considering the known life history attributes of P. torreyana, and the disturbance histories of the stands, predictions are made concerning the spatial characteristics of the respective stands. All trees within each study plot were sized by diameter breast height (DBH) and mapped. Three techniques of spatial analysis are applied to the resulting unweighted point pattern distributions and the distributions weighted by the square root of DBH as a surrogate for age. The results are consistent with predictions and confirm the following generalizations concerning patterns of aggregation in Torrey pine stands. Young trees tend to be more aggregated than old trees within the same stands. Young stands tend to be more aggregated than old stands on otherwise ecologically similar sites. On a periodically disturbed site there are clusters of trees that represent cohorts of post-disturbance recruitment.     

Comparison of life history traits between invasive and native populations of purple loosestrife (Lythrum salicaria) using nonlinear mixed effects model

We compared growth patterns of invasive and native populations of purple loosestrife (Lythrum salicaria) while varying water and nutrient levels. We examined three life-history traits (height, number of branches, and the size of largest leaf) during the growth period adopting a nonlinear mixed effects model. Invasive populations were found to be slower in shoot elongation but grew to be taller than native populations. Invasive populations produced more branches than natives only in the high water, high nutrient treatment. Invasive populations had a similar increase in the size of the largest leaf compared to natives, but ultimately produced a greater size of largest leaf than natives. Invasive populations were found to display a greater vegetative expansion, but this was not strongly affected by our treatments.     

Effects of increasing temperatures on population dynamics of the zebra mussel <Emphasis Type=Italic>Dreissena polymorpha</Emphasis>: implications from an individual-based model

Zebra mussels (Dreissena polymorpha, Pallas, 1771) have had unprecedented success in colonizing European and North American waters under strongly differing temperature regimes. Thus, the mussel is an excellent model of a species which is able to cope with increasing water temperatures expected under global change. We study three principle scenarios for successful survival of the mussel under rising temperatures: (1) no adaptation to future thermal conditions is needed, existing performance is great enough; (2) a shift (adaptation) towards higher temperatures is required; or (3) a broadening of the range of tolerated temperatures (adaptation) is needed. We developed a stochastic individual-based model which describes the demographic growth of D. polymorpha to determine which of the alternative scenarios might enable future survival. It is a day-degree model which is determined by ambient water temperature. Daily temperatures are generated based on long-term data of the River Rhine. Predictions under temperature conditions as recently observed for this river that are made for the phenology of reproduction, the age distribution and the shell length distribution conform with field observations. Our simulations show that temporal patterns in the life cycle of the mussel will be altered under rising temperatures. In all scenarios spawning started earlier in the year and the total reproductive output of a population was dominated by the events later in the spawning period. For maximum temperatures between 20 and 26°C no thermal adaptation of the mussel is required. No extinctions and stable age distributions over generations were observed in scenario 2 for all maximum temperatures studied. In contrast, no population with a fixed range of tolerated temperatures survived in scenario 3 with high maximum temperatures (28, 30, 32°C). Age distributions showed an excess of 0+ individuals which resulted in an extinction of the population for several thermal ranges investigated. Priority programme of the German Research Foundation—contribution 14.     

The Chapman-Enskog procedure for an age-structured population model: initial, boundary and corner layer corrections

We consider a mathematical model of an age-structured population of some fisheries (for example, anchovies, sardines or soles). Two time scales are involved in the problem: the fast time scale for the migration dynamics and the slow time scale for the demographic process. At a first step, we study the so called 'aggregated' system by means of the semigroups theory. Then, we study the asymptotic behaviour of the model by using the Chapman-Enskog procedure. In particular, we study initial, boundary and corner layer effects in order to obtain the exact initial and boundary conditions the approximated solution has to satisfy.     

The Paton individual-based model legacy

Ray Paton oversaw the creation of a long lineage of Individual-based Models (IbMs) and this paper discusses the five most successful. All of these concern the development of adaptation, covering both evolutionary time and organism lifetime (somatic time). Of the five models discussed here, the first is based on a plant-herbivore model, the other four are based on a substrate-bacteria model, with the option of antibiotics.     

A spatial simulation model for vegetation dynamics

A fine-resolution, spatially explicit, stochastic model was developed to simulate the dynamics of species cover abundance and pattern in a single vegetation layer wherein neighbouring individuals are assumed to compete for growing space. Each species in the model is characterized by a small number of morphological and life-history parameters, which enter into equations that stand for a minimal set of vegetation processes. The model performed well in reproducing post-fire successional trends among the three codominant dwarf shrubs in a Dutch heathland community as recorded in an annually mapped permanent quadrat. Program inputs, outputs and an example of sensitivity analysis are illustrated. With suitable changes, the model could potentially describe any plant community in which the vertical structure is simple and community dynamics are determined by spatial interactions among neighbouring plants.Jacques de Smidt provided data, information and stimulus for this project. The model was developed during ICP's visit to Utrecht, arranged by Marinus Werger and financed by The Netherlands Science Research Council (ZWO). We also thank David Glenn-Lewin and Ernst Lippe for discussion and cooperation.     

A semiparametric model for the analysis of recurrent-event panel data

In many longitudinal studies, interest focuses on the occurrence rate of some phenomenon for the subjects in the study. When the phenomenon is nonterminating and possibly recurring, the result is a recurrent-event data set. Examples include epileptic seizures and recurrent cancers. When the recurring event is detectable only by an expensive or invasive examination, only the number of events occurring between follow-up times may be available. This article presents a semiparametric model for such data, based on a multiplicative intensity model paired with a fully flexible nonparametric baseline intensity function. A random subject-specific effect is included in the intensity model to account for the overdispersion frequently displayed in count data. Estimators are determined from quasi-likelihood estimating functions. Because only first- and second-moment assumptions are required for quasi-likelihood, the method is more robust than those based on the specification of a full parametric likelihood. Consistency of the estimators depends only on the assumption of the proportional intensity model. The semiparametric estimators are shown to be highly efficient compared with the usual parametric estimators. As with semiparametric methods in survival analysis, the method provides useful diagnostics for specific parametric models, including a quasi-score statistic for testing specific baseline intensity functions. The techniques are used to analyze cancer recurrences and a pheromone-based mating disruption experiment in moths. A simulation study confirms that, for many practical situations, the estimators possess appropriate small-sample characteristics.     

The onion model, a simple neutral model for the evolution of diversity in bacterial biofilms

Bacterial biofilms are particularly resistant to a wide variety of antimicrobial compounds. Their persistence in the face of antibiotic therapies causes significant problems in the treatment of infectious diseases. Seldom have evolutionary processes like genetic drift and mutation been invoked to explain how resistance to antibiotics emerges in biofilms, and we lack a simple and tractable model for the genetic and phenotypic diversification that occurs in bacterial biofilms. Here, we introduce the 'onion model', a simple neutral evolutionary model for phenotypic diversification in biofilms. We explore its properties and show that the model produces patterns of diversity that are qualitatively similar to observed patterns of phenotypic diversity in biofilms. We suggest that models like our onion model, which explicitly invoke evolutionary process, are key to understanding biofilm resistance to bactericidal and bacteriostatic agents. Elevated phenotypic variance provides an insurance effect that increases the likelihood that some proportion of the population will be resistant to imposed selective agents and may thus enhance persistence of the biofilm. Accounting for evolutionary change in biofilms will improve our ability to understand and counter diseases that are caused by biofilm persistence.     

The identification of enzyme targets for the optimization of a valine producing Corynebacterium glutamicum strain using a kinetic model

The enzyme targets for the rational optimization of a Corynebacterium glutamicum strain constructed for valine production are identified by analyzing the control of flux in the valine/leucine pathway. The control analysis is based on measurements of the intracellular metabolite concentrations and on a kinetic model of the reactions in the investigated pathway. Data‐driven and model‐based methods are used and evaluated against each other. The approach taken gives a quantitative evaluation of the flux control and it is demonstrated how the understanding of flux control is used to reach specific recommendations for strain optimization. The flux control coefficients (FCCs) with respect to the valine excretion rate were calculated, and it was found that the control is distributed mainly between the acetohydroxyacid synthase enzyme (FCC = 0.32), the branched chain amino acid transaminase (FCC = 0.27), and the exporting translocase (FCC = 0.43). The availability of the precursor pyruvate has substantial influence on the valine flux, whereas the cometabolites are less important as demonstrated by the calculation of the respective response coefficients. The model is further used to make in‐silico predictions of the change in valine flux following a change in enzyme level. A doubling of the enzyme level of valine translocase will result in an increase in valine flux of 31%. By optimizing the enzyme levels with respect to valine flux it was found that the valine flux can be increased by a factor 2.5 when the optimal enzyme levels are implemented. © 2009 American Institute of Chemical Engineers Biotechnol. Prog., 2009     

The roles of predation,competition, and exploitation in the trophic dynamics of a warmwater stream: a model synthesis,analysis, and application

We developed a trophic dynamic model of key populations and processes in the New River, West Virginia, to identify the mechanisms most responsible for maintaining food web structure. Key populations were represented by thirteen model components and were aquatic insects; age-1 and age-2 crayfish (three species); age-1 and age-2 hellgrammites (Corydalus cornutus larvae); non-game fishes; age-0, age-1, and adult smallmouth bass (Micropterus dolomieu); age-0, age-1, and adult rock bass (Ambloplites rupestris); and age-0, age-1 to age-3, and adult flathead catfish (Pylodictis olivaris). In this system, crayfish and hellgrammites are harvested to provide bait for the recreational fishery that extensively exploits the three predatory fish species. Predation and intraspecific regulation were represented with nonlinear algorithms, and linear terms represented fishery harvests. Interspecific competition among components occurred through predation on shared prey. Error analysis of the model suggested that predation was the most important mechanism in maintaining system structure (the disposition of biomass among system components). Further, the trophic relation between each component and its prey accounted for 34–64% of the variability in food web structure, whereas predation on each component explained 1–24% of food web structure variability. Therefore, so-called ‘bottom-up’ effects were more influential than ‘top-down’ effects. Interspecific competition and intraspecific regulation had secondary roles in maintaining New River food web structure, although intraspecific regulation was most important to aquatic insects, which were not predatory in our model. Both forms of competition are probably tempered by extensive predation and exploitation in the New River system. Exploitation was a secondary structuring agent to adult smallmouth bass, which experience a high rate of harvest in the New River.