Abstract
Maintenance is the energy that living organisms are bound to use to maintain their structure in a viable state. It includes all the metabolic and physiological costs that are not directly associated to the production of biomass (growth and reproduction) or to development (maturation). In the framework of the DEB theory, somatic maintenance rate can either be proportional to organism structural volume V or, more marginally, to structural surface V 2/3. Being mostly associated to similar metabolic processes, volume specific maintenance costs are not expected to vary substantially at both intra-and interspecific levels. In the DEB theory, the volume-specific maintenance rate is therefore supposed to keep constant from birth to death and to remain approximately constant between species. However, a recent meta-analysis of DEB parameters estimated using the Add-my-Pet collection (Kooijman, 2014) reveals troubling patterns apparently violating this inter-specific scaling rule and challenging the DEB theory. It is indeed shown in this study that empirically-derived volume-specific maintenance rates scale approximately with, and display a very high variability around this trend. Overall, estimated maintenance rates in Add-my-Pet span over three to four orders of magnitude, thus invalidating the assumption of constant maintenance rate between species, which underpins the covariation rules for parameter values of the DEB theory. In an attempt to address this major problem for the DEB theory, we propose a simple physiological mechanism that would simultaneously explain the apparent decrease of volume-specific maintenance rate with ultimate size and its apparent variability for a given range of maximum size. Our proposition consists in making protein (and more generally structure) turnover explicit in maintenance and linking protein damage rate to aerobic metabolism and the production of ROS, which are decreasing with both structural volume and maximum structural volume. We show that this implies that the actual volume specific maintenance rate varies both at the intra- and inter-specific levels in a range very similar to what is observed in the Add-my Pet data estimations. If true, this implies that the apparent decrease of volume-specific maintenance rate with ultimate size is an artefact and it requires modifications of the standard DEB theory in order to capture empirical inter-specific scaling patterns of DEB parameters while keeping the consistency of the theory at both intra- and inter-specific levels.