Metabolism provides a basis for using first principles of physics, chemistry, and biology to link the biology of individual organisms to the ecology of populations, communities, and ecosystems. Metabolic rate, the rate at which organisms take up, transform, and expend energy and materials, is the most fundamental biological rate. We have developed a quantitative theory for how metabolic rate varies with body size and temperature. Metabolic theory predicts how metabolic rate, by setting the rates of resource uptake from the environment and resource allocation to survival, growth, and reproduction, controls ecological processes at all levels of organization from individuals to the biosphere. Examples include: (1) life history attributes, including devel- opment rate, mortality rate, age at maturity, life span, and population growth rate; (2) population interactions, including carrying capacity, rates of competition and predation, and patterns of species diversity; and (3) ecosystem processes, including rates of biomass production and respiration and patterns of trophic dynamics. Data compiled from the ecological literature strongly support the theoretical predictions. Even- tually, metabolic theory may provide a conceptual foundation for much of ecology, just as genetic theory provides a foundation for much of evolutionary biology.

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Toward a metabolic theory of ecology

Aconstitutive model is proposed for the deformation of rubber materials which is shown to represent successfully the response of these materials in uniaxial extension, biaxial extension, uniaxial compression, plane strain compression and pure shear. The developed constitutive relation is based on an eight chain representation of the underlying macromolecular network structure of the rubber and the non-Gaussian behavior of the individual chains in the proposed network. The eight chain model accurately captures the cooperative nature of network deformation while requiring only two material parameters, an initial modulus and a limiting chain extensibility. Since these two parameters are mechanistically linked to the physics of molecular chain orientation involved in the deformation of rubber, the proposed model represents a simple and accurate constitutive model of rubber deformation. The chain extension in this network model reduces to a function of the root-mean-square of the principal applied stretches as a result of effectively sampling eight orientations of principal stretch space. The results of the proposed eight chain model as well as those of several prominent models are compared with experimental data of Treloar (1944, Trans. Faraday Soc. 40, 59) illustrating the superiority, simplicity and predictive ability of the proposed model. Additionally, a new set of experiments which captures the state of deformation dependence of rubber is described and conducted on three rubber materials. The eight chain model is found to model and predict accurately the behavior of the three tested materials further confirming its superiority and effectiveness over earlier models.

https://hal.archives-ouvertes.fr/hal-01390807/document

A three-dimensional constitutive model for the large stretch behavior of rubber elastic materials

In this review I show that the '3/4-power scaling law' of metabolic rate is not universal, either within or among animal species. Significant variation in the scaling of metabolic rate with body mass is described mainly for animals, but also for unicells and plants. Much of this variation, which can be related to taxonomic, physiological, and/or environmental differences, is not adequately explained by existing theoretical models, which are also reviewed. As a result, synthetic explanatory schemes based on multiple boundary constraints and on the scaling of multiple energy-using processes are advocated. It is also stressed that a complete understanding of metabolic scaling will require the identification of both proximate (functional) and ultimate (evolutionary) causes. Four major types of intraspecific metabolic scaling with body mass are recognized [based on the power function R=aMb, where R is respiration (metabolic) rate, a is a constant, M is body mass, and b is the scaling exponent]: Type I: linear, negatively allometric (b 1) to one or two later phases of negative allometry (b<1). Ontogenetic changes in the metabolic intensity of four component processes (i.e. growth, reproduction, locomotion, and heat production) appear to be important in these different patterns of metabolic scaling. These changes may, in turn, be shaped by age (size)-specific patterns of mortality. In addition, major differences in interspecific metabolic scaling are described, especially with respect to mode of temperature regulation, body-size range, and activity level. A 'metabolic-level boundaries hypothesis' focusing on two major constraints (surface-area limits on resource/waste exchange processes and mass/volume limits on power production) can explain much, but not all of this variation. My analysis indicates that further empirical and theoretical work is needed to understand fully the physiological and ecological bases for the considerable variation in metabolic scaling that is observed both within and among species. Recommended approaches for doing this are discussed. I conclude that the scaling of metabolism is not the simple result of a physical law, but rather appears to be the more complex result of diverse adaptations evolved in the context of both physico-chemical and ecological constraints.

Beyond the '3/4-power law': variation in the intra- and interspecific scaling of metabolic rate in animals.

http://academic.oup.com/auk/article-pdf/102/3/661/30080634/auk0661.pdf

Scaling: why is animal size so important?

We examine the scaling law $B \propto M^{\alpha}$ which connects organismal metabolic rate $B$ with organismal mass $M$, where $\alpha$ is commonly held to be 3/4. Since simple dimensional analysis suggests $\alpha=2/3$, we consider this to be a null hypothesis testable by empirical studies. We re-analyze data sets for mammals and birds compiled by Heusner, Bennett and Harvey, Bartels, Hemmingsen, Brody, and Kleiber, and find little evidence for rejecting $\alpha=2/3$ in favor of $\alpha=3/4$. For mammals, we find a possible breakdown in scaling for larger masses reflected in a systematic increase in $\alpha$. We also review theoretical justifications of $\alpha=3/4$ based on dimensional analysis, nutrient-supply networks, and four-dimensional biology. We find that present theories for $\alpha=3/4$ require assumptions that render them unconvincing for rejecting the null hypothesis that $\alpha=2/3$.

https://arxiv.org/pdf/physics/0007096.pdf

Re-examination of the "3/4-law" of Metabolism

Scaling laws governing the cardiovascular system of mammals are discussed in the present review in a manner emphasizing their experimental basis. Specific attention is given to the well-known experimental laws requiring the rate of oxygen consumption and the heart rate of mammals to vary with body mass raised to the powers 3/4 and −1/4, respectively. This review involves reconsideration and further discussion of the previous work of the writer in which these and other scaling relationships were developed from fundamental considerations. The predicted scaling laws remain unchanged from the earlier work, but alternative assumptions leading to the laws are used so as to provide additional insight. The scaling laws are shown to have their origin in the basic design of the cardiovascular system and in the basic processes involved in its working. Modification of the design assumptions of the system to account for known differences in the relative heart masses of mammals and birds is shown to lead to the scaling laws for rate of oxygen consumption and heart rate of birds.

Similitude in the cardiovascular system of mammals.

Allometric Relations and Scaling Laws for the Cardiovascular System of Mammals

A general derivation is presented for the scaling laws governing the size and number of capillary blood vessels in mammals. The derivation is based on the assumption of three idealized similarity principles known to apply, at least approximately, to resting mammals: (i) size-invariant blood pressure; (ii) size-invariant fraction of blood in the capillaries; and (iii) size-invariant oxygen consumption and uptake, per unit of body mass, during each heart cycle. Results indicate that the radius and length of capillaries, and the number that are open and active in the resting state, should scale with mammal mass to the powers 1/12, 5/24 and 5/8, respectively, consistent with earlier work by the author. Measurements are presented supporting the results. Physiological changes accompanying strenuous exercise are accounted for by a change in the scaling law for capillary number, from scaling exponent 5/8 to 3/4.

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Scaling laws for capillary vessels of mammals at rest and in exercise

Vascular networks refer here mainly to the microscale capillary networks of the vascular system of mammals, although they may also be considered to include the small arteries that feed the capillaries and the small veins that drain them. The modeling of these networks for resting mammals is reviewed within the context of describing related scaling laws for mammals of vastly different size. Basic processes are considered and alternative approaches mentioned. All lead to the same scaling laws for the radius, length and number of the vessels. The applicability of the relations is illustrated using existing measurements. Discussion is also included on the effect of strenuous exercise on the scaling law for number of capillary vessels and matters related to it.

/pdf/modeling-of-vascular-networks-338ykr5whx.pdf

Modeling of vascular networks

In order to illustrate the theory of plasticity as developed in the previous chapter, we now proceed to consider a few selected problems requiring plasticity considerations for solution. Because of the inherently nonlinear form of the governing equations in plasticity theory, and the attendant mathematical complexities, these problems will be restricted to rather simple ones. Nevertheless they should serve to illustrate the general features involved in the plastic response of solids.

Thomas H. Dawson

Papers

Similitude in the cardiovascular system of mammals.

Allometric Relations and Scaling Laws for the Cardiovascular System of Mammals

Scaling laws for capillary vessels of mammals at rest and in exercise

Modeling of vascular networks

Problems in Plasticity