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Recently Caplan (1) applied the concepts of irreversible thermodynamics and cybernetics to contracting muscle and derived Hill's force-velocity relation. Wilkie and Woledge (2) then compared Caplan's theory to chemical rates inferred from heat data and concluded that the theory was not consistent with the data. Caplan defended his theory in later papers (3, 4) but without any direct experimental verifications. As Wilkie and Woledge (2) point out, the rate of phosphorylcreatine (PC) breakdown during steady states of shortening has not been observed because of technical difficulties. In this paper it is shown that the rate equations may be directly integrated with time to obtain relations among actual quantities instead of rates. The validity of this integration is based on experimental evidence which indicates that certain combinations of the transport coefficients are constant with muscle length. These equations are then directly compared to experimental data of Cain, Infante, and Davies (5) with the following conclusions: (a) The measured variations of DeltaPC for isotonic contractions are almost exactly as predicted by Caplan's theory. (b) The value of the chemical rate ratio, nu(m)/nu(o), obtained from these data was 3.53 which is close to the value of 3 suggested by Caplan (3). (c) The maximum value of the chemical affinity for PC splitting was found to be 10.6 k cal/mole which is as expected from in vitro measurements (2). Because of the excellent agreement between theory and experiment, we conclude that Caplan's theory definitely warrants further investigation.

A phenomenological theory for contracting muscle based on irreversible thermodynamics and the sliding filament theory is developed. The individual cross bridges, considered as subunits, are viewed as linear energy converters with constant transport coefficients. With this view of the subunits, phenomenological equations applicable to the whole muscle are obtained. The transport coefficients are shown to be a function of a single parameter which is the number of activated cross bridges at any instant. By requiring Hill's force-velocity relation (1) to be satisfied, the response of the muscle is related to the number of activated cross bridges. The resulting theory differs significantly from the theory developed by Caplan (2) and a comparison of the theories is presented. The theory is shown to correlate well with the heat data of Woledge (3) for a tortoise muscle and gives a value of Y (ratio of chemical affinity to enthalpy of reaction) equal to 0.945. The comparison of the theory with Hill's frog muscle data (1) and (4) is also encouraging. In part II of this series, length variations are considered and the resulting theoretical predictions are shown to be consistent with experimental data.

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In part I of this series, the theory of irreversible thermodynamics was applied to the sliding filament model to obtain rate equations for a contracting muscle at the in situ length l(o). In this paper we extend the theory to include length variations derived from the sliding filament model of contracting muscle using the work of Gordon, Huxley, and Julian (1). Accepting the validity of Hill's forcevelocity relation (2) at the in situ length, we show that Hill's equation is valid for any length provided that the values of the parameters, a, b, and V(m) vary with length as derived herein. The predicted variation with length of the velocity for a lightly loaded isotonic contraction is shown to agree well with that measured by Gordon, Huxley, and Julian (1). Chemical rates are derived as functions of length using parameters that can be obtained experimentally.