Energy transformation and entropy production in living systems I. Applications to embryonic growth.

Generalized material and energy balances are presented for biological systems that experience negligible kinetic, elastic, and potential energy changes. The balances are used to characterize the mass changes and energy transformations that occur in the developing avian embryo, using as an example a consistent set of data for the chicken egg. It is shown that the rate of total chemical energy turnover by the embryo is a quantity of interest and that this rate is not necessarily equivalent to the metabolic rate that is predicted from heat transfer measurements or oxygen consumption rates. The energy required for evaporative water loss is accounted for in the overall energy balance. Using the results of the energy calculations and a generalized expression for the rates of internal and total entropy production, the Prigogine-Wiame hypothesis is examined for the developing embryo with two different assumptions regarding the efficiency of biomass conversion. An order of magnitude analysis of the internal heat-conduction term is performed to show that the chemical reaction term dominates the entropy production relation. The constant efficiency case is shown to be in agreement with the Prigogine-Wiame hypothesis for the data used in the analysis.