@phdthesis{Patra2013,
  author    = {Patra, Pintu},
  title     = {Population dynamics of bacterial persistence},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-69253},
  school      = {Universit{\"a}t Potsdam},
  year      = {2013},
  abstract  = {The life of microorganisms is characterized by two main tasks, rapid growth under conditions permitting growth and survival under stressful conditions. The environments, in which microorganisms dwell, vary in space and time. The microorganisms innovate diverse strategies to readily adapt to the regularly fluctuating environments. Phenotypic heterogeneity is one such strategy, where an isogenic population splits into subpopulations that respond differently under identical environments. Bacterial persistence is a prime example of such phenotypic heterogeneity, whereby a population survives under an antibiotic attack, by keeping a fraction of population in a drug tolerant state, the persister state. Specifically, persister cells grow more slowly than normal cells under growth conditions, but survive longer under stress conditions such as the antibiotic administrations. Bacterial persistence is identified experimentally by examining the population survival upon an antibiotic treatment and the population resuscitation in a growth medium. The underlying population dynamics is explained with a two state model for reversible phenotype switching in a cell within the population. We study this existing model with a new theoretical approach and present analytical expressions for the time scale observed in population growth and resuscitation, that can be easily used to extract underlying model parameters of bacterial persistence. In addition, we recapitulate previously known results on the evolution of such structured population under periodically fluctuating environment using our simple approximation method. Using our analysis, we determine model parameters for Staphylococcus aureus population under several antibiotics and interpret the outcome of cross-drug treatment. Next, we consider the expansion of a population exhibiting phenotype switching in a spatially structured environment consisting of two growth permitting patches separated by an antibiotic patch. The dynamic interplay of growth, death and migration of cells in different patches leads to distinct regimes in population propagation speed as a function of migration rate. We map out the region in parameter space of phenotype switching and migration rate to observe the condition under which persistence is beneficial. Furthermore, we present an extended model that allows mutation from the two phenotypic states to a resistant state. We find that the presence of persister cells may enhance the probability of resistant mutation in a population. Using this model, we explain the experimental results showing the emergence of antibiotic resistance in a Staphylococcus aureus population upon tobramycin treatment. In summary, we identify several roles of bacterial persistence, such as help in spatial expansion, development of multidrug tolerance and emergence of antibiotic resistance. Our study provides a theoretical perspective on the dynamics of bacterial persistence in different environmental conditions. These results can be utilized to design further experiments, and to develop novel strategies to eradicate persistent infections.},
  language  = {en}
}
@misc{SiblyGrimmMartinetal.2013,
  author    = {Sibly, Richard M. and Grimm, Volker and Martin, Benjamin T. and Johnston, Alice S. A. and Kulakowska, Katarzyna and Topping, Christopher J. and Calow, Peter and Nabe-Nielsen, Jacob and Thorbek, Pernille and DeAngelis, Donald L.},
  title     = {Representing the acquisition and use of energy by individuals in agent-based models of animal populations},
  series = {Methods in ecology and evolution : an official journal of the British Ecological Society},
  volume    = {4},
  journal   = {Methods in ecology and evolution : an official journal of the British Ecological Society},
  number    = {2},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {2041-210X},
  doi       = {10.1111/2041-210x.12002},
  pages     = {151 -- 161},
  year      = {2013},
  abstract  = {Agent-based models (ABMs) are widely used to predict how populations respond to changing environments. As the availability of food varies in space and time, individuals should have their own energy budgets, but there is no consensus as to how these should be modelled. Here, we use knowledge of physiological ecology to identify major issues confronting the modeller and to make recommendations about how energy budgets for use in ABMs should be constructed. Our proposal is that modelled animals forage as necessary to supply their energy needs for maintenance, growth and reproduction. If there is sufficient energy intake, an animal allocates the energy obtained in the order: maintenance, growth, reproduction, energy storage, until its energy stores reach an optimal level. If there is a shortfall, the priorities for maintenance and growth/reproduction remain the same until reserves fall to a critical threshold below which all are allocated to maintenance. Rates of ingestion and allocation depend on body mass and temperature. We make suggestions for how each of these processes should be modelled mathematically. Mortality rates vary with body mass and temperature according to known relationships, and these can be used to obtain estimates of background mortality rate. If parameter values cannot be obtained directly, then values may provisionally be obtained by parameter borrowing, pattern-oriented modelling, artificial evolution or from allometric equations. The development of ABMs incorporating individual energy budgets is essential for realistic modelling of populations affected by food availability. Such ABMs are already being used to guide conservation planning of nature reserves and shell fisheries, to assess environmental impacts of building proposals including wind farms and highways and to assess the effects on nontarget organisms of chemicals for the control of agricultural pests.},
  language  = {en}
}
@article{MassieRyabovBlasiusetal.2013,
  author    = {Massie, Thomas Michael and Ryabov, Alexei and Blasius, Bernd and Weithoff, Guntram and Gaedke, Ursula},
  title     = {Complex transient dynamics of stage-structured populations in response to environmental changes},
  series = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences},
  volume    = {182},
  journal   = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences},
  number    = {1},
  publisher = {Univ. of Chicago Press},
  address   = {Chicago},
  issn      = {0003-0147},
  doi       = {10.1086/670590},
  pages     = {103 -- 119},
  year      = {2013},
  abstract  = {Stage structures of populations can have a profound influence on their dynamics. However, not much is known about the transient dynamics that follow a disturbance in such systems. Here we combined chemostat experiments with dynamical modeling to study the response of the phytoplankton species Chlorella vulgaris to press perturbations. From an initially stable steady state, we altered either the concentration or dilution rate of a growth-limiting resource. This disturbance induced a complex transient response-characterized by the possible onset of oscillations-before population numbers relaxed to a new steady state. Thus, cell numbers could initially change in the opposite direction of the long-term change. We present quantitative indexes to characterize the transients and to show that the dynamic response is dependent on the degree of synchronization among life stages, which itself depends on the state of the population before perturbation. That is, we show how identical future steady states can be approached via different transients depending on the initial population structure. Our experimental results are supported by a size-structured model that accounts for interplay between cell-cycle and population-level processes and that includes resource-dependent variability in cell size. Our results should be relevant to other populations with a stage structure including organisms of higher order.},
  language  = {en}
}
@phdthesis{Martin2013,
  author    = {Martin, Benjamin},
  title     = {Linking individual-based models and dynamic energy budget theory : lessons for ecology and ecotoxicology},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-67001},
  school      = {Universit{\"a}t Potsdam},
  year      = {2013},
  abstract  = {In the context of ecological risk assessment of chemicals, individual-based population models hold great potential to increase the ecological realism of current regulatory risk assessment procedures. However, developing and parameterizing such models is time-consuming and often ad hoc. Using standardized, tested submodels of individual organisms would make individual-based modelling more efficient and coherent. In this thesis, I explored whether Dynamic Energy Budget (DEB) theory is suitable for being used as a standard submodel in individual-based models, both for ecological risk assessment and theoretical population ecology. First, I developed a generic implementation of DEB theory in an individual-based modeling (IBM) context: DEB-IBM. Using the DEB-IBM framework I tested the ability of the DEB theory to predict population-level dynamics from the properties of individuals. We used Daphnia magna as a model species, where data at the individual level was available to parameterize the model, and population-level predictions were compared against independent data from controlled population experiments. We found that DEB theory successfully predicted population growth rates and peak densities of experimental Daphnia populations in multiple experimental settings, but failed to capture the decline phase, when the available food per Daphnia was low. Further assumptions on food-dependent mortality of juveniles were needed to capture the population dynamics after the initial population peak. The resulting model then predicted, without further calibration, characteristic switches between small- and large-amplitude cycles, which have been observed for Daphnia. We conclude that cross-level tests help detecting gaps in current individual-level theories and ultimately will lead to theory development and the establishment of a generic basis for individual-based models and ecology. In addition to theoretical explorations, we tested the potential of DEB theory combined with IBMs to extrapolate effects of chemical stress from the individual to population level. For this we used information at the individual level on the effect of 3,4-dichloroanailine on Daphnia. The individual data suggested direct effects on reproduction but no significant effects on growth. Assuming such direct effects on reproduction, the model was able to accurately predict the population response to increasing concentrations of 3,4-dichloroaniline. We conclude that DEB theory combined with IBMs holds great potential for standardized ecological risk assessment based on ecological models.},
  language  = {en}
}
@article{MartinJagerNisbetetal.2013,
  author    = {Martin, Benjamin T. and Jager, Tjalling and Nisbet, Roger M. and Preuss, Thomas G. and Grimm, Volker},
  title     = {Predicting population dynamics from the properties of individuals - a cross-level test of dynamic energy budget theory},
  series = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences},
  volume    = {181},
  journal   = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences},
  number    = {4},
  publisher = {Univ. of Chicago Press},
  address   = {Chicago},
  issn      = {0003-0147},
  doi       = {10.1086/669904},
  pages     = {506 -- 519},
  year      = {2013},
  abstract  = {Individual-based models (IBMs) are increasingly used to link the dynamics of individuals to higher levels of biological organization. Still, many IBMs are data hungry, species specific, and time-consuming to develop and analyze. Many of these issues would be resolved by using general theories of individual dynamics as the basis for IBMs. While such theories have frequently been examined at the individual level, few cross-level tests exist that also try to predict population dynamics. Here we performed a cross-level test of dynamic energy budget (DEB) theory by parameterizing an individual-based model using individual-level data of the water flea, Daphnia magna, and comparing the emerging population dynamics to independent data from population experiments. We found that DEB theory successfully predicted population growth rates and peak densities but failed to capture the decline phase. Further assumptions on food-dependent mortality of juveniles were needed to capture the population dynamics after the initial population peak. The resulting model then predicted, without further calibration, characteristic switches between small-and large-amplitude cycles, which have been observed for Daphnia. We conclude that cross-level tests help detect gaps in current individual-level theories and ultimately will lead to theory development and the establishment of a generic basis for individual-based models and ecology.},
  language  = {en}
}