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Thermodynamic Direction of Biological Evolution: Model and Reality


HomepageINSTITUTE of Physico-Chemical Problems of EvolutionBiological Evolution and AgingThermodynamic Direction of Biological Evolution: Model and Reality

G. P. Gladyshev

Biology Bulletin, Vol. 23, No. 4, 1996, pp. 315-322. Translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya, No. 4, 1996, pp. 389-397.

Original Russian Text Copyright © 1996 by Gladyshev.

Semenov Institute of Chemical Physics, Russian Academy of Sciences, ul. Kosygina 4, Moscow, 117977 Russia

Received May 28, 1995

Abstract—A macrothermodynamic model of evolution of the supramolecular structures and chemical composition of living objects during ontogenesis and at long-term stages of general biological evolution is presented. A study of quasiclosed (thermodynamically and kinetically) systems, phases of the biomass supramolecular structures, enables one to draw a conclusion on the thermodynamic direction of biological evolution. In correspondence with the second principle, this direction leads to variations in the chemical composition and structure of the living systems during their development. Indirect and direct evidence of the trend to a minimal Gibbs specific function of formation of the supramolecular structures in animal tissues during ontogenesis are presented. The conclusion that thermodynamics is the "driving force" of evolution of the biological world is confirmed.

"One of the principal objects of theoretical research in any department of knowledge is to find the point of view from which the subject appears in its greatest simplicity."
J. Wllard Gibbs (1881)

"In addition to entropy there may well exist other "one-way" functions which add to the overall description of the world as temporal development."
Kenneth G. Denbigh (1989b)

INTRODUCTION

The study of surrounding objects is a main task in science. Any study is based on models that are precise only by definition, since the real systems are more complicated and cannot be described within the framework of simple concepts. Of course, the models can be made more precise. But there is a certain limit to their improvement related to expediency and, if you like, common sense.

Understanding of evolution and behavior of the natural systems is to a great extent based on classical natural science. Two approaches have played a special role: thermodynamic and kinetic. Thermodynamic description of the systems and phenomena is based on the concept of equilibrium. Thermodynamics answers the question: where is the process directed before the equilibrium can be achieved? Thermodynamics does not operate the time as a parameter and does not consider mechanisms of the phenomena. On the contrary, kinetics studies the rates of processes and their mechanisms.

Foundations of nonequilibrium thermodynamics have been developed during recent decades for the systems close to equilibrium (irreversible processes). Nonequilibrium thermodynamics united both of the approaches mentioned. However the results obtained are so far applicable only to certain phenomena. Something similar can also be said about the thermodynamic systems remote from equilibrium or synergetics. They are both based on pure kinetic methods.

Limitations of the mentioned approaches led to the opinion that the evolution of living systems can hardly agree with the second principle of thermodynamics.

A new discipline has recently appeared, hierarchic thermodynamics or macrothermodynamics (Gladyshev, 1978, 1988, 1993, 1994, 1995b), which allows a study of living objects on the basis of equilibrium thermodynamics and physical chemistry of the natural systems (Calvin, 1971; Ponnamperuma, 1972; Fox and Dose, 1972; Tanford, 1994). Macrothermodynamics is also based on the principles of macrokinetics. In a sense, it is an alternative to thermodynamics of the systems close to equilibrium. Macrothermodynamic models can be used for studying weakly nonequilibrium processes of morphogenesis, which are analogs of phase transitions.

Approaches to development of the macrothermodynamic theory of biological evolution have already been outlined (Gladyshev, 1978, 1988). A model was developed, some elements of which were substantiated. At the same time, the model (as applied to physicochemical phenomena), has never been presented in a clear and concise form accessible to biologists. Undoubtedly, this complicates the understanding of the approach proposed by the authors and, mainly, does not allow for evaluation of the predicting force of the theory. This situation appears to be due to the author's aspiration to reveal the driving force of evolution only from general deductive positions, above all on the basis of reductionism. Besides, the absence of strict experimental evidence on model correctness prevented its wide application.

After the first publications (Gladyshev, 1978), substantial progress was achieved. It became possible to apply the model to evolution of the chemical composition of living objects on the basis of quantitative data and present evidence for correspondence between the model and reality (Gladyshev, 1987, 1988, 1994, 1995a).

Here I briefly describe a physicochemical model of a particular case of evolution of the living system: evolution of supramolecular structures (their chemical composition and structure), with special reference to the main assumptions of the model. In addition, some experimental results are provided, which confirm the model applicability.

MODEL OF EVOLUTION OF THE LIVING SYSTEM

Let us consider a given volume of biological tissue (biomass) as a heterogeneous thermodynamic system consisting of a. liquid phase, aqueous solution of physiological substances, and a phase of supramolecular structures (supramolecular "skeleton" of biological structures), which appeared as a result of aggregation (self-assembly) of the molecules and supramolecular structures of various hierarchies (Gladyshev, 1994, 1995a). The phase of supramolecular structures appears as a result of weakly nonequilibrium phase transitions (to what extent this concept corresponds to the classical concept of phase is a matter for further discussion).

It is assumed that local supramolecular equilibrium is established in all points (microvolumes) of the phase of supramolecular structures, where small molecules are also present. Hence, we will call self-assembly leading to formation of the phase of supramolecular structures thermodynamic self-organization, unlike dynamic self-organization or simply self-organization (Prigogine, 1980) observed in the systems remote from equilibrium. The existence of local equilibrium means that we deal with a community of thermodynamically quasiclosed microvolumes, components of the considered phase (at times comparable with the duration of establishment of this equilibria). Hence, it is evident that the integral value of specific (averaged by volume) Gibbs (or Helmholtz) function of formation of the "averaged local conformation" of supramolecular structures achieves a minimum:

, (1)

For the phase of supramolecular structures of constant composition (at times of establishment of local equilibrium).

where V is the system volume, m is the mass of micro volumes, x, y, and z are coordinates; symbol "-" means that is a specific value; and symbol "~" underlines the heterogenous pattern of the system. Note that expression (1) appears not to be questioned.

Naturally, the biological system is open at times markedly exceeding the times of establishment of local intermolecular equilibria: a flow of substance passes through it. The system is as if swollen and its total volume and mass increase. The model implies that the mean flow of matter is stationary (the flow velocity oscillates around its mean value) and the nature of matter incoming to the system (phase of supramolecular structures) remains practically unchanged. In other words, the supramolecular phase (structure) of the organism is evolved "against the background" of the incoming flow of chemical substances (of practically constant composition). If the flow is sufficiently slow, it can be assumed that the liquid phase of the biological system is always in equilibrium with the flow and this provides, on average, for constant concentrations of the incoming substances at this phase, which, therefore, can be considered together with the environment as a thermostat, in a wide sense of the term, for the phase of supramolecular structures.

The assumption about constant and averaged in time flow of chemical substances into the biological system from the environment has been experimentally substantiated. It was shown (Gladyshev, 1993, 1994) that sequences of natural hierarchic structures (according to the energies of their formation) correspond to those of mean life (relaxation) times of these structures, e.g., it can be written for an individual community of certain closely related organisms

(I)

where t is the mean life time of "free" molecules-metabolites, supramolecular structures, organelles, cells in the biological tissue, organisms, populations, or communities.

Sequence (I) is a geometrical progression of the type , where is the mean life time of structures of the nth hierarchy in a certain biological system; n-l,2,3...,n; is the standard time equal to the mean life time for the structure of a lower (standard) hierarchy (0) of the sequence in question; b is the constant for the given sequence.

The law (I) allows us to distinguish between the thermostat (environment) and the system studied j per se, which forms a complete thermodynamic system [j + (j + 1)] together with its thermostat (j + 1). As was already mentioned, this suggests the existence of quasiclosed systems in the biological world, which function against the background of practically constant kinetic factors that determine the flow of substances from the environment, i.e., thermostat, and makes it possible to avoid, to a certain extent, insurmountable obstacles of using functions of the state for description of behavior of the open systems of this type.

Thus, the mean life time of individual cells of the organism (or its organ) is, as a rule, many factors of ten less than the life span of the organism itself. Hence, it is clear that the medium of the organism (organ) is a thermostat for the component cells. The presence of the thermostat allows us, as will be mentioned below, to consider the cell or their community as a kinetic quasiclosed system.

The chemical composition of the phase of supramolecular structures of the biological system slowly changes at times comparable with the duration of adaptive processes and ontogenesis, as well as during phylogenesis and at long-term stages of biological evolution as a whole. With the biological tissue senescence, the supramolecular structures become more thermodynamically stable (the supramolecular structures themselves, rather than the chemical substances that form these structures).

Selection of thermodynamically more stable suprastructures (structural stabilization of the phase) is determined by the thermodynamic factor: it is assumed that the time of retention (term taken from chromatography) of molecules (macromolecules) in the supramolecular phase is connected with the Gibbs function of formation of the supramolecular structures:

(2)

where R is the gas constant.

The molecules retained in the supramolecular medium for the longest period of time (incoming to the biological system from the environment or products of biosynthesis) enhance selection of similar molecules and this also changes the composition (and chemical nature) of the phase of supramolecular structures. As was already mentioned, this change is due to the thermodynamic factor, although expressed through kinetics (2). Thus, molecules are accumulated in the phase of supramolecular structures, whose absorption (self-assembly) is most thermodynamically profitable (these molecules have increased affinity to the phase of supramolecular structures). With the mechanisms of template synthesis, these molecules have advantages during reduplication (reproduction). As a result, specific Gibbs function of formation of the superstructures (or specific Helmholtz function practically coinciding with the former in the condensed phase) increases in the absolute value during evolution of the biological tissue and becomes more negative. Hence, it follows that

(3)

For the phase of supramolecular structure of variable composition (at times of ontogenesis, phylogenesis, etc.)

Expression (3) means that the value of (1) attaining a minimum at local equilibrium () gradually changes during ontogenesis (phylogenesis and at long-term stages of evolution) tending to an even lower value ().

Let us note for clarity that equation (3) is a consequence of partial kinetic quasicloseness of the phase of supramolecular structures (or biological tissue as a whole) for outgoing flows of the matter. This quasicloseness leads to accumulation of supramolecular structures of the aggregates of molecules with increased thermodynamic stability in a single volume (Gladyshev, 1988, pp. 90-92). Equations (2) and (3) set, in fact, the axis of time (kinetic parameter) for alteration of the values and and, hence, their sum, whose value is below zero:

(4)

However the last conclusion (as was already mentioned) refers to the case of flow of the matter (constant in composition) incoming to the biological system. If the composition of the incoming flow changes in time, e. g„ in case of certain physiological abnormalities, the kinetic quasicloseness of the system can be disturbed and the pattern of change becomes indeterminate.

Fig.1. Schematic diagram of changes in specific values, components of the Gibbs function of biological system j during ontogenesis and phylogenesis. The scheme is also applicable to long-term stages of biological evolution Biological system, j, for example, animal tissue. and , specific values of chemical (ch) and supramolecular (im) component of the jth system; and scales of time of ontogenesis and phylogenesis, respectively; scales and without scale ; functions , , , are set arbitrarily: values of a with indices characterize specific values of the components of the Gibbs function for structure formation (specific free Gibbs energy of formation of the corresponding structure) of the system at a specific moment of time; values of a in the course of evolution change as a result of changes in the chemical composition of organisms or species.

The thermodynamically open adsorption (absorption) system, through which the flow of substances of constant composition gradually enters, which are subject to phase or chemical transformations in this system, is a simple analog for the presented model (Gladyshev, 1988, pp. 90-92). Thus, if fatty acid and water are the main components of a homogenous flow at the input, and microemulsion, in which the fatty acid concentration is high, is formed in the column (reactor), the column is rather rapidly "filled" with fatty acid (partial kinetic quasicloseness of the system according to the outgoing flow of the matter is thus expressed) and ceases to function. In this case, the trend presented by expression (3) cannot, apparently, be questioned (Gladyshev, 1988, pp. 71-75, 90-92, 163-165).

Expression (3) means that during senescence, the normal biological tissue should be enriched with chemical compounds having the most negative Gibbs function of the formation of suprastructures. Chemically energy consuming substances with relatively least negative Gibbs functions of the formation of chemical compounds, , such as lipids, proteins, polysaccharides, and nucleic acids, which force water from the biological tissue in the course of its aging, are such compounds, as follows from the theory and were experimentally confirmed (Stull et al., 1969; Gladyshev, 1988; Gladyshev and Kitaeva, 1995). Such trends should be observed during phylogenesis and at long-term stages of biological evolution, when the chemical composition of the environment should be assumed as constant (in this case, the biological systems may be considered as partially kinetically quasiclosed).

The model considers self-assembly irrespective of the regime (equilibrium or nonequilibrium) of chemical reactions in the liquid phase. Formation of the supramolecular phase moves to the background of the mechanisms of synthesis of the substances and their transfer: these substances are "used" for construction of the supramolecular structure of the biological tissue. The model also implies that the considered thermodynamic system is simple: by definition, only the work of extension can be realized in it (this work is relatively small in the condensed phase). Of course, this approximation becomes unjustified when studying, for example, evolution of the population, a structure of high hierarchic level (where the organisms per se play the role of interacting particles and irreversible phenomena not accompanied by changes of entropy are studied), which performs mechanical or other work (Gladyshev, 1988, 1995b; Denbigh, 1989b).

Functioning of the biological systems, e.g., biological tissue, is possible on the condition of their sufficient "permeability" for the matter-building material of the supramolecular structure. Besides, not only internal, but also external forces should be present that enhance "mixability" inside the system, metabolism (Gladyshev, 1994). Periodic oscillations of the environmental parameters (thermostat) around the mean values play the role of such forces. Let us stress that the essential periodically changing external factors are an inseparable "thermodynamic effect" of the environment on evolution of the biological system. Hence, the joint effects of internal thermodynamic factors (expressed inside the system) and external thermodynamic effects (changes and oscillations of the environmental physical parameters) determine the direction of evolution.

This model pays special attention to physical chemistry of the supramolecular structures, which should be considered as a "key" for understanding biological evolution. It can be easily proved (Gladyshev, 1988) that the model does not contradict the kinetic theory of Darwin and Walles and reconciles it with many critics of this theory.

EXPERIMENTAL EVIDENCE OF THE MODEL

In accordance with the presented model, the biological system is swollen under the normal physiological conditions during ontogenesis and, according to equations (3) and (4), it is enriched with energy-consuming chemical substances, which force water from the biological tissue. Figure 1 presents a theoretical scheme of changes in the supramolecular (on) and chemical (ch) components of the specific Gibbs function of the biological tissue during ontogenesis (ont) and phylogenesis (ph). In previous publications, the chemical component of Gibbs function was designated as m (molecule). This scheme, (Gladyshev, 1978) now receives convincing experimental confirmation.

Indeed, extensive materials have been accumulated on changes in the overall chemical composition of the organs and tissues during ontogenesis, phylogenesis, and biological evolution as a whole. Figure 2 presents a characteristic example of variation of the chemical composition (water-organic substances) of the animals brain as a function of its relative evolutionary development. These data (Gladyshev, 1978) were taken from the reference literature. A comparatively high confidence interval for humans (point 1 in Fig.2), just as in all other cases (points 2-14, Fig.2; confidence intervals were not indicated for all points), is due to variation in the chemical composition of the brain tissue during ontogenesis; the results are usually given without indication of the age of animals. It follows from Fig.2 that during evolution, the brain of animals, as was already mentioned, is enriched with fats, proteins, and other chemically energy-consuming organic compounds. Now the task consists in showing that the discussed variation of the chemical composition is a sequence of the trend of the evolving biological system to the supramolecular equilibrium. In other words, equations (3) and (4) should be experimentally confirmed.

One way to do this is related to the selection of a physicochemical parameter of the phase of supramolecular structure or individual elements of this phase, which would enable the estimation of the formation of this phase (characterizing the thermostability of structures) and chemical energy capacity of the component substances. The temperature of melting , of supramolecular structures of the corresponding condensed phase can be such a parameter.

Fig. 2. The brain content of water as a function of its relative development for the .jth animal () (Gladyshev, 1977). It is assumed that is the linear function of the brain content of water. For humans , and for frogs . The mean data are given for adult individuals: (1) humans; (2) monkey; (3) horse; (4) dog; (5) cat; (6) rabbit; (7) duck; (8) mouse: (9) guinea pig; (10) rat; (11) shark, carp and gull; (12) peach; (13) turtle; (14) frog.

Let us begin with theoretical concepts. For the closed systems, the relationship of Gibbs function and T is determined by the Gibbs-Helmholtz equation:

, (5)

where D H is a change of enthalpy, T and p are the temperature and pressure. A similar equation can also be written for the systems (p and V are constant), whose composition is not identical. This assumption will be precise only for the ith substances, which have similar values of , and , where index refers to the point of melting of the ith substances. Nevertheless, this assumption proved to be reasonable (in the case of variation in thermodynamic characteristics of the substances in a certain range), since a correlation was found between , specific Gibbs function of non equilibrium phase transition "supercooled liquid-solid body" (at the standard temperature 298 K) and for a wide range of organic compounds with changing in a wide interval of temperatures (Gladyshev, 1988, pp. 161-163; Vakulichev and Gladyshev, 1989; Gladyshev and Gladyshev, 1994). Calculations can be performed using our approximated equation, an analogue of the approximated Gibbs-Helmholtz equation:

, (6)

where and the upper index im indicates that the substance condensation is considered.

Some experimental results on fatty acids are presented in Fig. 3. It can be seen that the studied natural substances with elevated tend to form more thermodynamically stable phases, solid structures (more negative ), since processes of condensation (crystallization) of pure substances can be considered as enthalpy-controlled (from the viewpoint of total effect). For more precise calculations, corrections should be introduced for a change in heat capacity of the substances during their melting, (see equation (8)). Note that the chemical energy capacity of some monotypic natural compounds () increases with their . This confirms the pattern of predicted changes in during ontogenesis and evolution as a whole (Fig. 1).

In the case of entropy-controlled processes, e. g., cell aggregation during self-assembly of cell structures, denaturation of some proteins, etc., the increase in thermostability of the structures is accompanied by a decrease in their T„ . However it can be shown that the relative proportion of the above-mentioned entropy-controlled processes is comparatively small during formation of the structure of some biological tissues or organelles.

Fig. 3. Specific Gibbs function of nonequilibrium phase transition "supercooled liquid-solid body" as a function of at 298 K for some fatty

acids; and is specific Gibbs function of crystallization (solidification) and temperature of melting of the ith compound. The value of is calculated per unit mass. Correlation is preserved when is calculated per unit volume. (1) Saturated fatty acids; (2) unsaturated fatty acids.

Thus, the data in Fig. 3 confirm the conclusion that the living organisms thermodynamically economize from accumulation of chemically high energy-consuming substances, which force water from the biological tissue. The model was convincingly confirmed by the adaptive thermodynamically directed change in the composition of fatty acids involved in the synthesis of fats (Gladyshev, 1995a).

Many data suggest that the cells of microorganisms, plants, and animals change in composition of fatty acids (fats) with changes in the environmental temperature. At the body temperature below the optimum, the content of unsaturated fatty acids (fatty acid residues) with a lower melting temperature () (as compared with the saturated and other unsaturated acids) increases. This increase is easily detected from the growth of the iodine number of the corresponding fatty acid fractions. At the temperature above the optimum, the proportion of saturated acids with a high temperature of melting increases. The results of a series of studies are given elsewhere (Aleksandrov, 1975; Gladyshev, 1995a) and in the reference literature.

If we do not touch upon kinetic problems, i.e., do not consider the mechanisms responsible for compensatory changes in the composition of fatty acids, but consider the phenomenon from the position of macrothermodynamics of formation of the lipid structures (against the background of constant kinetic factors, i.e., on the condition of the above mentioned kinetic quasicloseness of the system), the following conclusion can be drawn.

These patterns can be easily understood on the basis of the macrotbennodynamic model on assuming that the ratio of enthalpy-control to entropy-control during formation of condensed "solid" monotypic structures containing fatty acids undergoes insignificant changes in the adaptive interval of temperatures.

On the basis of the data presented in Fig. 3, it can be easily shown that the observed changes in the chemical composition of fatty acids (fats) of the biological tissues during ontogenesis are thermodynamically profitable upon changes in temperature.

According to the thermodynamic model of adaptation, a flow of the matter (of constant composition) enters the unit volume V of fat tissue (cell) from a thermostat (liquid phase or surrounding medium), in which a constant concentration of fatty acids (fats) is maintained. This volume is similar to a chromatographic column, where the incoming substances are absorbed. An evident relationship can be used for calculation:

, (7)

where and are concentrations of fatty acids in the volume V in the solid aggregated (adsorbed) state and in the liquid phase, respectively. is the change in the specific Gibbs function during adsorption (self-assembly) of fatty acids.

For example, when the temperature changes from 25°C to 5°C, the amount of palmitic, elaidic and oleic (b ) acids in the tissue will increase by a factor of 1.07, 1.23,1.53, respectively, according to the model calculations (Gladyshev, 1995a). This fully agrees with the above mentioned facts indicating that the relative amount of unsaturated acids increases and that of saturated acids decreases with the decrease of body temperature.

The above is also illustrated by the classical data of Frenkel and Hopp on dependence of the iodine number of phosphatides (lipids) of the fly larvae on the temperature of their rearing (Aleksandrov, 1975). At an increased temperature, the amount of unsaturated acids decreases. These data agree with the results of model calculations: at a decrease of temperature by 10°C, the iodine number increases by a factor of 1.21 ± 0.03.

Thus, the calculations confirm applicability of the macrotherroodynamic model to evolution of the fat tissue. Variations in the chemical composition of fatty acids (fats) during adaptation to the environmental temperature can be explained within the framework of this model.

Some results pertaining to variations in the chemical composition of proteins and nucleic acids in evolution have already been discussed (Gladyshev, 1988; Schuster, 1993). However, there is so far insufficient information, which would unambiguously confirm the thermodynaroic direction of changes in the composition of these natural polymers. The increase in the temperature of chromatin melting during ontogenesis is the most demonstrative example. This fact suggests the thermodynamic nature of evolutionary aging during ontogenesis (Gladyshev, 1988, pp. 161-163). New results (Schuster, 1993) suggest that evolutionary optimization of the RNA structure is determined by the thermodynamic stability of not only its secondary structure, but also of its tertiary structure. This explains the fact why not only sequences with GC pairs are selected during evolution, which is, on the whole, most profitable from the viewpoint of thermodynamics of the secondary structure formation. Sequences with AU pairs arc also selected during evolution. Hence, thermodynamics of the tertiary and other higher supramolecular structures affects the RNA chemical composition and structure, as follows from the theory. Thus, I believe that selection of natural (AUGC) sequences is most profitable from the viewpoint of macrothermodynamics. Schuster considers these sequences as most stable with respect to mutations.

Data have recently been published, which enable unambiguous calculations suggesting the thermodynamic direction of the development of animal tissue during ontogenesis*(*- Additional information, References). For example, the influence of the age of the tail tendon collagen tissue on the temperature and heat of its denaturation was studied by differential scanning calorimetry in rats (Tsereteli and Belopol'skaya, 1994). Examination of approximately 100 samples has shown that when the age of this tissue changes from two weeks to two years, the temperature of its denaturation (melting) increases by approximately 6°C: from 55°C to 64.5°C. According to our estimate, the enthalpy of denaturation changes from 6.0 to 7.6 Cal/g. The heat capacity changes during transition of the tissue from the native state to the denaturated one:

. (8)

On the basis of the presented data from the analog of the Gibbs-Helmholtz equation taking into account a change of heat capacity during phase transition, we can easily and with sufficient precision (without accounting for variation of heat capacity of the system with an increase of temperature) calculate a change in the Gibbs function during formation of the tissue supramolecular structure at the standard (reference) temperature, :

. (9)

Assuming that T = 298.2 K (25°C) for the tissue of a 2-year old individual, we will have:

(10)

And for the tissue of a 2-week old animal

(11)

Calculations for various stages of ontogenesis suggest that in accordance with the theory, the value of of intact collagen of the rat tail tendon tends to a minimum during aging and changes in the above-mentioned case to

(12)

Note that the rough estimate according to the approximated equation

(13)

gives . Changes of (and ; Fig. 1) during the collagen tissue aging agiee quite well with variations of the water content, which changes in the range of 78-58% weight (Nildtin et al., 1977).

The values of obtained by us for the collagen tissue are less than similar data for foimation of the condensed (solid) phase of some chemical compounds (Gladyshev, 1995a; Gladyshev and Kitaeva, 1995). This does not contradict the physical picture of the phenomena of life, the principle of semistable state, and other known facts (Nildtin et al., 1977; Denbigh, 1968; Tanford, 1994).

There are grounds to believe that these experimental results may be very important and will stimulate studies with the aim of obtaining further direct evidence of applicability of the second principle to evolution of the biological and other natural systems. General methods are required for cognition of the world. It looks like the method of macrothermodynamics is one such method.

CONCLUSION

The available experimental data suggest that the macrothermodynamic model of evolution of living systems is applicable to the real world: the thermodynamic force is the driving force of biological evolution.

The first work on thermodynamics of biological evolution was published 20 years ago and I would like to recall that this work was published thanks to the decision of the late Editor-in-Chief of the Journal of Theoretical Biology, Dr. James Danielli, F.R.S., the author of the first model of biological membrane, an outstanding theoretical biologist and physicochemist of our century. The manuscript of this work was reviewed eight times and, as a result, the Editor-in-Chief wrote a letter to the author of this paper. It said, specifically: "Today I decided to publish your paper, although the western reviewers experience serious difficulties in its estimation. I took this decision because this work may be outstanding". Now the results of direct experiments have confirmed the correctness of the macrothermodynamic model of biological evolution. Dr. Danielli was not mistaken in his decision. There are grounds to believe that the method of macrothermodynamics (despite limitations) can be sufficiently effective in studies of ontogenesis, phylogenesis, and biological evolution as a whole on a solid physical basis.

REFERENCES

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  20. Stull, D.R„ Westnim, E.FJr., Sinke, G.C., The Chemical Thermodynamics of Organic Compounds, NewYoric: Wiley, 1969, Tanford, Ch., The Hydrophobic Effect and the Organization of Living Matter, Origins of Life. The Central Concepts.
  21. Deamer, D.W. and Fleischaker, G.R., Eds., Boston: Jones and Bartlett, 1994, pp. 233-239.
  22. Tsereteli, G.I. and Belopol'skaya, TV., Biofizika, 1994, vol. 39, no. 5, pp. 793-794.
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* - Additional information in:

  1. Structural and thermodynamic proporties of DNA uncover different evolutionary histories.
    Miramontes P, Medrano L, Cerpa C, Cedergren R, Ferbeyre G, Cocho G
    J Mol Evol 1995 Jun 40:6 698-704
  2. Towards protein folding by global energy optimization.
    Abagyan RA
    FEBS Lett 1993 Jun 28 325:1-2 17-22
  3. Structural and thermodynamic analysis of compensating mutations within the core of chicken egg white Iysozyme.
    Wilson KP, Malcolm BA, Matthews BW
    J Biol Chem 1992 May 25 267:15 10842-9
  4. Molecular evolution of plants as deduced from changes in free energy of 5S ribosomal RNAs.
    Mashkova TD, Barciszewska MZ, Joachimiak A, Nalaskowska M, Barciszewski J
    Int J Biol Macromol 1990 Aug 12:4 247-50
  5. A differential scanning calorimetry analysis of the age-related changes in the thermal stability of rat skin collagen.
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    Biochim Biophys Acta 1984 Dec 7 791:2 205-211
  6. See also: http://biomednet.com/db/medline
    Thermodynamics and Evolution, Thermodynamics and Aging,...

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