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Item The combined effects of reactant kinetics and enzyme stability explain the temperature dependence of metabolic rates(John Wiley and Sons Ltd, 2017-04-23) DeLong, J.P.; Gibert, J.P.; Luhring, Thomas M.; Bachman, G.; Reed, B.; Neyer, A.; Montooth, K.L.A mechanistic understanding of the response of metabolic rate to temperature is essential for understanding thermal ecology and metabolic adaptation. Although the Arrhenius equation has been used to describe the effects of temperature on reaction rates and metabolic traits, it does not adequately describe two aspects of the thermal performance curve (TPC) for metabolic rate—that metabolic rate is a unimodal function of temperature often with maximal values in the biologically relevant temperature range and that activation energies are temperature dependent. We show that the temperature dependence of metabolic rate in ectotherms is well described by an enzyme-assisted Arrhenius (EAAR) model that accounts for the temperature-dependent contribution of enzymes to decreasing the activation energy required for reactions to occur. The model is mechanistically derived using the thermodynamic rules that govern protein stability. We contrast our model with other unimodal functions that also can be used to describe the temperature dependence of metabolic rate to show how the EAAR model provides an important advance over previous work. We fit the EAAR model to metabolic rate data for a variety of taxa to demonstrate the model's utility in describing metabolic rate TPCs while revealing significant differences in thermodynamic properties across species and acclimation temperatures. Our model advances our ability to understand the metabolic and ecological consequences of increases in the mean and variance of temperature associated with global climate change. In addition, the model suggests avenues by which organisms can acclimate and adapt to changing thermal environments. Furthermore, the parameters in the EAAR model generate links between organismal level performance and underlying molecular processes that can be tested for in future work. © 2017 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.Item Size-dependent predation and correlated life history traits alter eco-evolutionary dynamics and selection for faster individual growth(Springer Tokyo, 2018-03-05) DeLong, J.P.; Luhring, Thomas M.Age at maturation is a key life history trait influencing individual fitness, population age structure, and ecological interactions. We investigated the evolution of age at maturity through changes in the von Bertalanffy growth constant for organisms with a simple juvenile-adult life history. We used Gillespie eco-evolutionary models to uncover the role of predation in driving the evolution of the growth constant when eco-evolutionary dynamics are present. We incorporated both size-independent and size-dependent predation into our models to generate differences in selection and dynamics in the system. Our results generally support the idea that faster ontogenetic growth is beneficial when populations are growing but that predation tends to have little effect on age at maturity unless there are trade-offs with other life history traits. In particular, if faster ontogenetic growth comes at the cost of fecundity, our results suggest that predation selects for intermediate levels of growth and fecundity. Eco-evolutionary dynamics influenced the nature of selection only when growth was linked to fecundity. We also found that predators that increasingly consume larger prey tend to have higher population sizes due to the greater energy intake from larger prey, but the growth rate-fecundity trade-off reversed this pattern. Overall, our results suggest an important role for interactions between size-dependent foraging and life-history trade-offs in generating varying selection on age at maturity through underlying growth traits. © 2018, The Society of Population Ecology and Springer Japan KK, part of Springer Nature.Item Predators modify the temperature dependence of life-history trade-offs(John Wiley and Sons Ltd, 2018-09) Luhring, Thomas M.; Vavra, Janna M.; Cressler, Clayton E.; DeLong, J.P.Although life histories are shaped by temperature and predation, their joint influence on the interdependence of life-history traits is poorly understood. Shifts in one life-history trait often necessitate shifts in another-structured in some cases by trade-offs-leading to differing life-history strategies among environments. The offspring size-number trade-off connects three traits whereby a constant reproductive allocation (R) constrains how the number (O) and size (S) of offspring change. Increasing temperature and size-independent predation decrease size at and time to reproduction which can lower R through reduced time for resource accrual or size-constrained fecundity. We investigated how O, S, and R in a clonal population of Daphnia magna change across their first three clutches with temperature and size-independent predation risk. Early in ontogeny, increased temperature moved O and S along a trade-off curve (constant R) toward fewer larger offspring. Later in ontogeny, increased temperature reduced R in the no-predator treatment through disproportionate decreases in O relative to S. In the predation treatment, R likewise decreased at warmer temperatures but to a lesser degree and more readily traded off S for O whereby the third clutch showed a constant allocation strategy of O versus S with decreasing R. Ontogenetic shifts in S and O rotated in a counterclockwise fashion as temperature increased and more drastically under risk of predation. These results show that predation risk can alter the temperature dependence of traits and their interactions through trade-offs. © 2018 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.Item Stoichiometry and life-history interact to determine the magnitude of cross-ecosystem element and biomass fluxes(Frontiers Media S.A., 2017-05-09) Luhring, Thomas M.; DeLong, J.P.; Semlitsch, R.D.Ecosystems are linked through the transfer of materials and energy. Studies examining material fluxes across habitat boundaries frequently quantify unidirectional flows of nutrients and energy. However, material fluxes can be multidirectional, and we lack a conceptual framework to describe how their quantity and stoichiometry influence the net transfer of individual elements between ecosystems. Here we develop a zero net transfer isocline (ZNTI) framework that integrates the relative mass and stoichiometry of fluxes into and out of an ecosystem. We then use case studies with amphibians and salmon to elucidate how life history, ontogenetic shifts in stoichiometry, and trophic interactions shape relative fluxes of nutrients between aquatic and terrestrial ecosystems. Because they increase in both size and Ca content from ova to metamorphs, amphibian life histories strongly bias them toward net Ca export into the terrestrial environment. Because amphibian biomass, C, P, and Ca ZNTIs do not overlap, there is no value of survivorship where the net flux of biomass, C, P, and Ca are simultaneously balanced between terrestrial and aquatic habitats. The degree of iteroparity and semelparity in salmon strongly affects both the magnitude of net biomass and P flux between riverine and marine environments. While the net direction of biomass flux generally remains strongly biased toward import into the riverine system, net P flux can reach net export into the marine environment because of increasing adult breeding survival leading to reduced mass and %P of what they deposit in rivers (e.g., ova vs. whole carcasses). These examples highlight how ontogenetic shifts in body size and stoichiometry result in asymmetric fluxes of elements and biomass that can lead to simultaneous net imports and exports of different elements within the same system. Furthermore, they demonstrate how changes in life-history characteristics and stage-specific survivorship can lead to changes in net elemental transport between ecosystems. © 2017 Luhring, DeLong and Semlitsch.Item A semelparous fish continues upstream migration when exposed to alarm cue, but adjusts movement speed and timing(Academic Press, 2016-11-01) Luhring, Thomas M.; Meckley, T.D.; Johnson, N.S.; Siefkes, M.J.; Hume, J.B.; Wagner, C.M.Animals make trade-offs between predation risk and pursuit of opportunities such as foraging and reproduction. Trade-offs between antipredator behaviours and foraging are well suited to manipulation in laboratory and field settings and have generated a vast compendium of knowledge. However, much less is known about how animals manage trade-offs between predation risk and pursuit of reproductive opportunities in the absence of the confounding effects of foraging. In the present study, we investigated how the nonfeeding migratory life stage of sea lamprey, Petromyzon marinus, responds to odour from dead conspecifics (a cue that induces avoidance behaviours in laboratory and field studies). We released groups of PIT-tagged sea lamprey 65m from the shore of Lake Michigan or 287m upstream in Carp Lake River and used antennas to detect their movements in the river. As the breeding season progressed, sea lamprey initiated upstream movement earlier and were more likely to enter the river. Sea lamprey that began the night in Lake Michigan entered Carp Lake River at higher rates and accelerated upstream when exposed to high concentrations of alarm cue, consistent with animals attempting to minimize time spent in risky areas. Sea lampreys that began the night in the river delayed upstream movement when exposed to alarm cue, consistent with animals sheltering and gathering information about a source of risk. We attribute this context-specific reaction to alarm cue to differences in perceived vulnerability to predation in sheltered positions in the river versus exposed positions in the lake. Once in the river, the vast majority of sea lamprey moved upstream independent of alarm cue or Julian date. Although life-history-induced time and energy budgets place rigid constraints on the direction of migration, sea lamprey attend to predation risk by modifying movement timing and speed. © 2016 The Association for the Study of Animal Behaviour