title: Negative relationships between population density and metabolic rates are not general.
abstract: Population density has recently been suggested to be an important factor influencing metabolic rates and to represent an important 'third axis' explaining variation beyond that explained by body mass and temperature. In situations where population density influences food consumption, the immediate effect on metabolism acting through specific dynamic action (SDA), and downregulation due to fasting over longer periods, is well understood. However, according to a recent review, previous studies suggest a more general effect of population density per se, even in the absence of such effects. It has been hypothesized that this results from animals performing anticipatory responses (i.e. reduced activity) to expected declines in food availability. Here, we test the generality of this finding by measuring density effects on metabolic rates in 10 clones from two different species of the zooplankton Daphnia (Daphnia pulex Leydig and D. magna Straus). Using fluorescence-based respirometry, we obtain high-precision measures of metabolism. We also identify additional studies on this topic that were not included in the previous review, compare the results and evaluate the potential for measurement bias in all previous studies. We demonstrate significant variation in mass-specific metabolism among clones within both species. However, we find no evidence for a negative relationship between population density and mass-specific metabolism. The previously reported pattern also disappeared when we extended the set of studies analysed. We discuss potential reasons for the discrepancy among studies, including two main sources of potential bias (microbial respiration and declining oxygen consumption due to reduced oxygen availability). Only one of the previous studies gives sufficient information to conclude the absence of such biases, and consistent with our results, no effect of density on metabolism was found. We conclude that population density per se does not have a general effect on mass-specific metabolic rate.
authors.affiliation: Centre for Biodiversity Dynamics, Department of Biology, NTNU, Norwegian University of Science and Technology, Realfagbygget, NO-7491, Trondheim, Norway.
authors.forename: Varvara
authors.initials: V
authors.lastname: Yashchenko
authors.affiliation: Centre for Biodiversity Dynamics, Department of Biology, NTNU, Norwegian University of Science and Technology, Realfagbygget, NO-7491, Trondheim, Norway.
authors.forename: Erlend Ignacio
authors.initials: EI
authors.lastname: Fossen
authors.affiliation: Centre for Biodiversity Dynamics, Department of Biology, NTNU, Norwegian University of Science and Technology, Realfagbygget, NO-7491, Trondheim, Norway.
authors.forename: Øystein Nordeide
authors.initials: ØN
authors.lastname: Kielland
authors.affiliation: Centre for Biodiversity Dynamics, Department of Biology, NTNU, Norwegian University of Science and Technology, Realfagbygget, NO-7491, Trondheim, Norway.
authors.forename: Sigurd
authors.initials: S
authors.lastname: Einum
mesh_terms: D000818:Animals; D003621:Daphnia / Q000502:physiology*; D004734:Energy Metabolism; D011156:Population Density; D013045:Species Specificity; D015048:Zooplankton / Q000502:physiology*
keywords: abundance; competition; digestion; genetic variation; metabolic theory of ecology; oxygen consumption; respiration