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Biomedical subjects

Van M Savage

Publications and source records attributed to Van M Savage.

9 recordsLinked to original sources

A quantitative, theoretical framework for understanding mammalian sleep.

Sleep is one of the most noticeable and widespread phenomena occurring in multicellular animals. Nevertheless, no consensus for a theory of its origins has emerged. In particular, no explicit, quantitative theory exists that elucidates or distinguishes between the myriad hypotheses proposed for sleep. Here, we develop a general, quantitative theory for mammalian sleep that relates many of its fundamental parameters to metabolic rate and body size. Several mechanisms suggested for the function of sleep can be placed in this framework, e.g., cellular repair of damage caused by metabolic processes as well as cortical reorganization to process sensory input. Our theory leads to predictions for sleep time, sleep cycle time, and rapid eye movement time as functions of body and brain mass, and it explains, for example, why mice sleep approximately 14 hours per day relative to the 3.5 hours per day that elephants sleep. Data for 96 species of mammals, spanning six orders of magnitude in body size, are consistent with these predictions and provide strong evidence that time scales for sleep are set by the brain's, not the whole-body, metabolic rate.

Animals↗

Kinetic effects of temperature on rates of genetic divergence and speciation.

Latitudinal gradients of biodiversity and macroevolutionary dynamics are prominent yet poorly understood. We derive a model that quantifies the role of kinetic energy in generating biodiversity. The model predicts that rates of genetic divergence and speciation are both governed by metabolic rate and therefore show the same exponential temperature dependence (activation energy of approximately 0.65 eV; 1 eV = 1.602 x 10(-19) J). Predictions are supported by global datasets from planktonic foraminifera for rates of DNA evolution and speciation spanning 30 million years. As predicted by the model, rates of speciation increase toward the tropics even after controlling for the greater ocean coverage at tropical latitudes. Our model and results indicate that individual metabolic rate is a primary determinant of evolutionary rates: approximately 10(13) J of energy flux per gram of tissue generates one substitution per nucleotide in the nuclear genome, and approximately 10(23) J of energy flux per population generates a new species of foraminifera.

Animals↗

Comment on "The illusion of invariant quantities in life histories".

Nee et al. (Reports, 19 August 2005, p. 1236) used a null model to argue that life history invariants are illusions. We show that their results are largely inconsequential for life history theory because the authors confound two definitions of invariance, and rigorous analysis of their null model demonstrates that it does not match observed data.

Analysis of Variance↗

The metabolic basis of whole-organism RNA and phosphorus content.

Understanding the storage, flux, and turnover of nutrients in organisms is important for quantifying contributions of biota to biogeochemical cycles. Here we present a model that predicts the storage of phosphorus-rich RNA and whole-body phosphorus content in eukaryotes based on the mass- and temperature-dependence of ATP production in mitochondria. Data from a broad assortment of eukaryotes support the model's two main predictions. First, whole-body RNA concentration is proportional to mitochondrial density and consequently scales with body mass to the -1/4 power. Second, whole-body phosphorus content declines with increasing body mass in eukaryotic unicells but approaches a relatively constant value in large multicellular animals because the fraction of phosphorus in RNA decreases relative to the fraction in other pools. Extension of the model shows that differences in the flux of RNA-associated phosphorus are due to the size dependencies of metabolic rate and RNA concentration. Thus, the model explicitly links two biological currencies at the individual level: energy in the form of ATP and materials in the form of phosphorus, both of which are critical to the functioning of ecosystems. The model provides a framework for linking attributes of individuals to the storage and flux of phosphorus in ecosystems.

Animals↗

Body sizes of hosts and parasitoids in individual feeding relationships.

In a natural community of 49 species (12 species of aphids and 37 species of their parasitoids), body lengths of 2,151 parasitoid individuals were, to an excellent approximation, related to the body lengths of their individual aphid hosts by a power law with an exponent close to 3/4. Two alternative models predict this exponent. One is based on surface area to volume relationships. The other is based on recent developments in metabolic ecology. Both models require a changing ratio (in both host and parasitoid) of length to diameter with increasing body length. These changing ratios are manifested differently in the two models and result in testably different predictions for the scaling of body form with increasing size. The estimated exponent of 3/4 for the relationship between individual host body size and individual parasitoid body size degrades to an exponent of nearly 1/2, and the scatter in the relationship between aphid and parasitoid body length is substantially increased, if the average length of a parasitoid species is examined as a function of the average length of its aphid host species instead of using measurements of individuals.

Animals↗

Improved approximations to scaling relationships for species, populations, and ecosystems across latitudinal and elevational gradients.

Historically, allometric equations relate organismal traits, such as metabolic rate, individual growth rate, and lifespan, to body mass. Similarly, Boltzmann or Q(10) factors are used to relate many organismal traits to body temperature. Allometric equations and Boltzmann factors are being applied increasingly to higher levels of biological organization in an attempt to describe aggregate properties of populations and ecosystems. They have been used previously for studies that analyse scaling relationships between populations and across latitudinal gradients. For these kinds of applications, it is crucial to be aware of the "fallacy of the averages", and it is often problematic or incorrect to simply substitute the average body mass or temperature for an entire population or ecosystem into allometric equations. We derive improved approximations to allometric equations and Boltzmann factors in terms of the central moments of body size and temperature, and we provide tests for the accuracy of these approximations. This framework is necessary for interpreting the predictions of scaling theories for large-scale systems and grants insight into which characteristics of a given distribution are important. These approximations and tests are applied to data for body size for several taxonomic groups, including groups with multiple species, and to data for temperature at locations of varying latitude, corresponding to ectothermic body temperatures. Based on these results, the accuracy and utility of these approximations as applied to biological systems are assessed. We conclude that approximations to allometric equations at the species level are extremely accurate. However, for systems with a large range in body size, evaluating the skewness and kurtosis is often necessary, so it may be advantageous to calculate the exact form for the averaged scaling relationships instead. Moreover, the improved approximation for the Boltzmann factor, which uses the average and standard deviation of temperature, is quite accurate and represents a significant improvement over previous approximations.

Animals↗

Effects of body size and temperature on population growth.

For at least 200 years, since the time of Malthus, population growth has been recognized as providing a critical link between the performance of individual organisms and the ecology and evolution of species. We present a theory that shows how the intrinsic rate of exponential population growth, rmax, and the carrying capacity, K, depend on individual metabolic rate and resource supply rate. To do this, we construct equations for the metabolic rates of entire populations by summing over individuals, and then we combine these population-level equations with Malthusian growth. Thus, the theory makes explicit the relationship between rates of resource supply in the environment and rates of production of new biomass and individuals. These individual-level and population-level processes are inextricably linked because metabolism sets both the demand for environmental resources and the resource allocation to survival, growth, and reproduction. We use the theory to make explicit how and why rmax exhibits its characteristic dependence on body size and temperature. Data for aerobic eukaryotes, including algae, protists, insects, zooplankton, fishes, and mammals, support these predicted scalings for rmax. The metabolic flux of energy and materials also dictates that the carrying capacity or equilibrium density of populations should decrease with increasing body size and increasing temperature. Finally, we argue that body mass and body temperature, through their effects on metabolic rate, can explain most of the variation in fecundity and mortality rates. Data for marine fishes in the field support these predictions for instantaneous rates of mortality. This theory links the rates of metabolism and resource use of individuals to life-history attributes and population dynamics for a broad assortment of organisms, from unicellular organisms to mammals.

Animals↗

Effects of size and temperature on developmental time.

Body size and temperature are the two most important variables affecting nearly all biological rates and times. The relationship of size and temperature to development is of particular interest, because during ontogeny size changes and temperature often varies. Here we derive a general model, based on first principles of allometry and biochemical kinetics, that predicts the time of ontogenetic development as a function of body mass and temperature. The model fits embryonic development times spanning a wide range of egg sizes and incubation temperatures for birds and aquatic ectotherms (fish, amphibians, aquatic insects and zooplankton). The model also describes nearly 75% of the variation in post-embryonic development among a diverse sample of zooplankton. The remaining variation is partially explained by stoichiometry, specifically the whole-body carbon to phosphorus ratio. Development in other animals at other life stages is also described by this model. These results suggest a general definition of biological time that is approximately invariant and common to all organisms.

Amphibians↗