What is Life from a Physical Chemist
On Thermodynamics, Entropy and Evolution of Biological
Systems: What is Life from a Physical Chemist's Viewpoint
Georgi P. Gladyshev
Abstract: Life in the Universe originated and is
evolving in accordance with the general laws of nature, specifically, the
law of temporal hierarchies and the second law of thermodynamics.
Key words: Second law of thermodynamics, Entropy, Law of temporal hierarchies,
Supramolecular thermodynamics, Biological Evolution.
"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 
Understanding of the evolution and behavior of natural systems is to
a great extent based on classical natural science. Two approaches have
played a special role: the thermodynamic and the kinetic. The thermodynamic
(thermostatic) description of systems and phenomena is based on the concept
of equilibrium. Thermodynamics answers the question: where is the process
directed before equilibrium can be achieved? Thermodynamics does not use
time as a parameter and does not consider the mechanism of the phenomenon.
Conversely, kinetics studies the rates of processes and their mechanisms.
The foundations of nonequilibrium thermodynamics have been developed
during recent decades for systems close to equilibrium (irreversible processes).
Nonequilibrium thermodynamics unites both equilibrium thermodynamics and
kinetics. However the results obtained are so far applicable only to certain
phenomena. The study of thermodynamic systems far from equilibrium, or
synergetics, is similarly limited. They are both based on pure kinetic
Limitations of these approaches led to the opinion that the evolution
of living systems cannot agree with the second principle of thermodynamics
A new discipline has recently appeared, hierarchic thermodynamics or
macrothermodynamics [3-11], which allows a study of living objects on the
basis of equilibrium thermodynamics and the physical chemistry of natural
systems [4,12,13]. Macrothermodynamics is also based on the principles
of macrokinetics. In a sense, it is an alternative to the thermodynamics
of systems close to equilibrium. Macrothermodynamic models can be used
for studying weakly nonequilibrium processes of structure formation, which
are analogs of phase transitions.
Here I briefly describe a physicochemical model of a particular case
of evolution of living systems: evolution of supramolecular structures
(their chemical composition and structure) with special reference to the
main assumptions of the model.
EVOLUTION, TEXTURE AND GENERAL LAWS
Evolution. In its common sense biological evolution is considered
an irreversible process of the historical variation of life with respect
to the evolutionary time scale characterizing the given object. So, Charles
Darwin and Alfred Wallace's theories and the modern synthetic theory of
evolution emphasize the evolution of populations. These theories study
the causes, mechanisms and general rules of the evolution of living organisms
from the biological point of view. Darwin's theory forms the theoretical
basis of all biology.
The development of modern darwinism is connected with the analysis of
the data obtained by molecular biology. It is aimed at a more profound
understanding of hereditary variability and at finding ways to control
living natural resources.
However, up to now the theory of biological evolution did not attempt
to point out the physical essence of evolutionary trends, although Darwin
admitted that "the principles of life are a part or a consequence of some
general law" determining the evolution of matter as a whole.
It becomes much easier to comprehend nature and the motive force of
biological evolution generally, as well as phylogenesis and ontogenesis,
if one examines the evolution of the composition and texture, the structure
of a biological object, and its hierarchic subsystems.
Texture is defined as the structure of the biological world.
Analyzing matter from the angle of its composition and texture, one is
struck by the hierarchic organization of the biological world. Thus, a
population is a sum total of organisms, which in turn consist of cells.
Cells consist of organelles and other complex supramolecular formations,
which themselves consist of macromolecules and low-molecular weight compounds.
Hence, the concept of a hierarchic thermodynamic system as a system consisting
of hierarchic subsystems can be introduced. There is every reason to believe
that any higher (j) hierarchic structure emerges as a result of
self-assembly, i.e. the thermodynamic self-organization of lower-hierarchy
(j-1) structures. It turns out that the average life-span of the
structures that constitute the elements of any lower hierarchy in the biomass
is significantly shorter than the average life-span of any higher-hierarchy
The law of temporal hierarchies. To exactly formulate the law
of temporal hierarchies (Gladyshev’s law), let us consider a biological
system consisting of the given organism’s cells, the organism itself, and
the population formed by these organisms (i.e., a fragment of the hierarchic
sequence of biological structures). Identifying the average life-span (life
time) of structures makes it possible to assert that the average life-span
(t) of a cell (cel) in the organism is much less than the
average life-span of the organism (org), which, in its turn, is
much less than the life-span of the population (pop):
This law can be laconically formulated as follows: structures of lower
hierarchy (j) live (exist) in biosystems with much shorter life-spans
than structures of higher hierarchy (j+1). This law, which can be
extended to all the hierarchies of the biological world, in a sense, "sets
the rules" for the formation of self-reproducing, identical, polyhierarchic
structures. Each higher structural hierarchy creates the habitat (the thermostat,
in the broad physical sense of R. Kubo , i.e. the habitat where at certain
intervals of time a number of parameters, including the concentrations
of chemical substances, are constant) for all lower-hierarchy structures.
If time hierarchies did not exist, the substances in such a world would
stay in the state of "homogeneous-heterogeneous mixture", and there simply
would be no phenomenon of life. The law of temporal hierarchies, which
correlates with the energy and spatial hierarchies of biological structures,
determines the existence of an exchange of chemical substances and the
organism’s other hierarchic structures in a living system.
The second law of thermodynamics is one of the general laws of
nature, which establishes the direction and the degree of completeness
of real thermodynamic processes. With regard to a nonequilibrium simple
isolated system such as a perfect gas (i.e., a system with constant internal
energy and volume in which no work is performed or only the work of expansion
is performed), the second law establishes that should some spontaneous
(irreversible) processes occur, a certain state function, called the system’s
entropy of R.Clausius and J.W.Gibbs, S, increases and tends
towards a maximum value.
For other types of systems, the second law is formulated by mathematically
equivalent definitions using other state functions. Thus, for simple closed
systems when temperature and pressure are constant, the criterion of the
spontaneity of processes is a change of the simple-system Gibbs function
(the Gibbs energy), G, which tends towards a minimum. For
simple closed systems where temperature and volume are constant, the criterion
of the processes’ spontaneity is a change of the simple-system Helmholtz
function, F, which also tends towards a minimum. As was pointed
out above, the operation of the law of temporal hierarchies in the biological
world makes it possible to consider each structural hierarchy of a real
open biological system as a thermodynamic system located in a thermostat
(i.e., an environment with constant parameters, including the intensity
of the sun's radiation). In such an open system, self-assembly processes
can be observed. At certain intervals of time, this system may be considered
as a quasi-closed system tending towards an increase of the thermodynamic
stability of its suprastructure, which is formed as a result of the interaction
(self-assembly) of the elementary structures comprising it. In this case,
if the temperature, pressure and other parameters of the thermostat are
constant, the second law determines the trend towards a minimum of the
specific value of the Gibbs function of the formation of the system in
Let us note again that depending on the type of thermodynamic system
identified, the second law in classical thermodynamics is formulated in
terms of the change of the various state functions. It is clear that formulating
this law of nature from the point of view of a change of entropy for real
systems is only a particular case and as such has limited significance.
Moreover, the thermodynamic criterion of the production of entropy characterizes
only the spontaneous heat processes in the simple system and doesn’t apply
for the non-entropy processes which take place in real complex systems
[1, 5]. Many authors forget this, thus causing confusion and serious misunderstandings.
As has recently been shown, the evolution of the composition and texture
of living organisms may be comprehended on the basis of the thermodynamic
study of different-level hierarchic structures. This makes it possible
to identify the motive force of evolution within the framework of the above-mentioned
general laws of nature without relying on the ideas of dynamic self-organization
(or just self-organization, using I. Prigogine’s terms) and dissipative
structures or on some supposedly still unknown "laws" of nature.
For example, the available data on the thermodynamic stability of supramolecular
biological structures and variations in the chemical composition of living
organisms have allowed a macrothermodynamic model of biological evolution
to be developed experimentally. In this model, the tendency toward a minimum
of the specific Gibbs function of the formation of supramolecular structures
of living organisms causes variations in the chemical composition and structure
of living systems. It is shown that during the course of ontogenesis and
phylogenesis, as well as long-lasting stages in the evolution of the organic
world, biosystems (as a result of the thermodynamic direction of evolutionary
processes of the formation of supramolecular structures) are enriched with
energy-intensive chemical substances, which displace water from these biosystems.
The change in the chemical composition and structure of biostructures of
an adaptive character is also explained from the angle of macrothermodynamics
The theory outlined offers an answer to questions about the origins
of life in the Universe and the motive forces of evolution and aging. The
answer is "the thermodynamic force".
MODEL OF EVOLUTION OF THE LIVING SYSTEMS
Let us consider a given volume of biological tissue (biomass) as a heterogeneous
thermodynamic system consisting of a liquid phase, an 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 [8,14]. The phase of supramolecular structures appears as a
result of weakly nonequilibrium phase transitions .
It is assumed that local supramolecular equilibrium is established at
all points (microvolumes) of the phase of supramolecular structures where
small molecules are also present [5,15]. Hence, we will call self-assembly
leading to formation of the phase of supramolecular structures thermodynamic
self-organization, unlike dynamic self-organization or simply Prigogine's
self-organization observed in systems far from equilibrium. The existence
of local equilibrium means that we deal with a community of thermodynamically
quasi-closed microvolumes, components of the considered phase (at times
comparable with the duration of establishment of this equilibrium). Hence,
it is evident that the integral value of the specific (averaged by volume)
Gibbs (or Helmholtz) function of formation of the "averaged local conformation"
of supramolecular (intermolecular) structures, achieves
For the phase of supramolecular structures of constant composition
(at times of relaxation to local equilibrium)
where V is the volume of the system;
m is the mass of the selected microvolumes;
x, y and
z are the coordinates; the sign "-" means that the specific
value is considered; and the sign "~" points
out the heterogeneous nature of the system. Note that expression (2) appears
to be generally accepted [13,15].
Naturally, a biological system is open at times markedly exceeding the
times of establishment of local intermolecular equilibrium: a flow of substances
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 quasi-stationary
(the flow velocity oscillates around its mean value) and the nature of
incoming matter to the system (phase of supramolecular structures) remains
practically unchanged. In other words, the supramolecular phase (structure)
of the organism evolves "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 in this phase, which,
therefore, can be considered together with the environment as a thermostat,
in the broad sense of the term, for the phase of supramolecular structures.
The assumption about constant and time-averaged flow of chemical substances
into the biological system from the environment has been experimentally
substantiated. It has been shown [5,14] that sequences of natural hierarchic
structures correspond to the series of average life-spans (life times)
of these structures in the biomass or biosystem. For instance, for a separate
community of several close species one can write:
where t is the mean life-span (life
time) of "free" metabolite molecules, supramolecular structures, organells,
cells, organisms, populations, and communities.
Sequence (3) is a geometrical progression of the type ,
the mean life-span of structures of the n-th
hierarchy in a certain biological system; n
- l,2,3..., n; is
the standard time equal to the mean life-span for the structure of a lower
(standard) hierarchy (0) of the sequence in question; and
is a constant for the given sequence.
The law (3) 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 already mentioned, this suggests the existence of quasi-closed
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 state
for the description of the behavior of open systems of this type.
Thus, the mean life-span of individual cells of the organism (or an
organ) is, as a rule, a few dozens times 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 the community
of cells as a kinetic quasi-closed 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 the long-term stages of biological evolution as a whole. With the senescence
of biological tissue, the supramolecular structures become more thermodynamically
stable (the supramolecular structures themselves, but not 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 (a term taken from chromatography)
of molecules (macromolecules) in the supramolecular phase,
is connected with the Gibbs function of formation of the supramolecular
where A is a coefficient that slowly
changes as the chemical composition of biological object changes in the
course of evolution (in principle each microvolume is described by its
coefficient A), and 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 photosynthesis and biosynthesis) enhance selection of similar
molecules, and this also changes the composition (and chemical nature)
of the phase of the supramolecular structures. As was already mentioned,
this change is due to the thermodynamic factor, although expressed through
kinetics (4). Thus, molecules accumulate in the phase of the supramolecular
structures whose absorption (self-assembly) is most thermodynamically profitable
(these molecules have higher affinity for the phase of the supramolecular
structures). If there are mechanisms of matrix synthesis, such molecules
have advantages during reduplication (reproduction). As a result, the specific
Gibbs function of formation of the superstructures, (or
specific Helmholtz function practically coinciding with the former in the
condensed phase), increases in absolute value during evolution of biological
tissues and becomes more negative. Hence, it follows that
For the phase of supramolecular structures of varying composition
(for the times of ontogenesis, phylogenesis, etc.)
Expression (5) means that the value of attaining
a minimum at local equilibrium ()
gradually changes during ontogenesis (and phylogenesis and at long-term
stages of evolution) tending to an even lower value ().
Let us note for clarity that equation (5) is a consequence of the partial
kinetic quasi-closeness of the phase of the supramolecular structures (or
biological tissue as a whole) for outgoing flows of matter. This
quasi-closeness leads to accumulation of supramolecular structures of the
aggregates of molecules with increased thermodynamic stability in a single
volume (microvolume). Equations (4) and (5) set, in fact, the axis of time
(kinetic parameter) for alteration of the values
and, hence, their sum, whose value is below zero:
This criterion (<0)was
used for the description the processes of evolution, aging and growth of
biological objects [3,5,9,11]
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
quasi-closeness of the system can be disturbed and the variation of becomes
The thermodynamically open system of adsorption (absorption) through
which the flow of substances of constant composition occurs, which are
subject to phase or chemical transformations in this system, is a simple
analog for the model presented . Thus, if fatty acid and water are the
main components of a homogenous flow at the input, and the microemulsion,
in which the fatty acid concentration is high, is formed in the column
(reactor), the column is rather rapidly "filled" with the fatty acid (partial
kinetic quasi-closeness of the system according to the outgoing flow of
matter is thus expressed) and ceases to function. In this case, the trend
presented by expression (5) cannot, apparently, be questioned .
Expression (5) means that during senescence, normal biological tissue
should be enriched with chemical compounds having the most negative
Gibbs function of the formation of suprastructures. Energy (chemical)
consuming substances with the relatively least negative (or more
positive) Gibbs function of the formation of chemical compounds, ,
such as lipids, proteins, polysaccharides, and nucleic acids, which force
water from biological tissue in the course of aging, are such compounds.
This follows from the thermodynamic theory of aging and has been experimentally
confirmed [5,9,17,18]. Such trends should be observed during phylogenesis
and at the long-term stages of biological evolution when the chemical composition
of the environment can be assumed constant (in this case, the biological
systems may be considered as partially kinetically quasi-closed).
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 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 a 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
Functioning of biological systems, e.g., biological tissue, is possible
on the condition of 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, e.g.,
metabolism. 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 the 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 the physical chemistry of the supramolecular
structures, which should be considered as a "key" for understanding biological
evolution. It can be easily proved  that the model does not contradict
the kinetic theory of Darwin and Wallace and reconciles it with many critics
of this theory.
EXPERIMENTAL EVIDENCE FOR THE MODEL
In accordance with the model presented, the biological system is swollen
under the normal physiological conditions during ontogenesis and, according
to equations (5) and (6), is enriched with energy-consuming (energy-intensive)
chemical substances, which force water from biological tissue. Previous
studies [3,5,11] present a theoretical scheme of changes in the supramolecular
(im) and chemical (ch) components of the specific Gibbs function
of biological tissue during ontogenesis (ont) and phylogenesis (ph).
This scheme has received convincing experimental confirmation.
Indeed, energy-intensive materials accumulate due to changes in the
overall chemical composition of the organs and tissues during ontogenesis,
phylogenesis, and biological evolution as a whole. The papers [3,11] present
a characteristic example of the variation of the chemical composition (water-organic
substances) of the brain as a function of relative evolutionary development.
It follows from these studies that during evolution, the brain of animals,
as was already mentioned, is enriched with fats, proteins, and other chemically
energy-consuming organic compounds. It has been shown that the variation
of the chemical composition is a consequence of the trend of the evolving
biological system to supramolecular equilibrium. In other words, equations
(5) and (6) are experimentally confirmed.
Data have been published, which enable unambiguous calculations suggesting
the thermodynamic direction of the development of animal tissues during
ontogenesis. For example, the influence of the age of rat skin collagen
on the temperature and heat of denaturation was studied by differential
scanning calorimetry (DSC) .
On the basis of the data presented from the Gibbs-Helmholtz equation
taking into account a change of heat capacity during the phase transition,
we can easily and with sufficient precision (without accounting for the
variation in the heat capacity of the system with an increase of temperature)
calculate a change in the Gibbs function during formation of the supramolecular
structure at the standard (reference) temperature.
Calculations for various stages of ontogenesis show that in accordance
with the theory, the value of of
intact collagen of the rat skin tends to a minimum during aging.
New results have also been obtained by the DSC method in the laboratory
of Professor J. Lepock. These data are in a good agreement with the theory
The principle of the stability of supramolecular structures requires
further elaboration. The biological systems that we have been discussing
are apparently governed by the principle of the stabilization of chemical
substances (Gladyshev's principle). The gist of this principle can be generally
stated as follows: during the formation of the more stable structures of
a higher hierarchical level (j), for example supramolecular structures,
nature spontaneously and predominantly uses the least stable structures
of a lower hierarchical level, i.e., the molecular level (j-1).
For instance, the molecular (chemically) stable substances such as H2,
N2, O2, CO2, H2O have relatively
low melting and boiling points which indicates qualitatively the low thermodynamic
stability of their condensed phases. On the other hand, energy-intensive
substances (with low molecular thermodynamic stability) such as sugars,
peptides, nucleic acids melt at relatively high temperatures and decompose
during melting and "boiling". The aggregated phases of these substances
are highly stable! Thus, the higher the chemical thermodynamic stability
of a substance, the lower its supramolecular thermodynamic stability in
the condensed state. The principle defined here is in agreement with the
experimental data and calculations done for the Gibbs - Helmholtz equation
It is very important to bear in mind that the principle of the stability
of chemical substances (and the general principle of the stability of substance
of other hierarchies) is a thermodynamic principle. Accordingly, the tendency
of biological systems during evolution (ontogenesis and phylogenesis) to
generate relatively highly stable structures of higher hierarchies leads
to the selection of relatively less stable structures of lower hierarchies.
That evolutionary tendency of the development of biological systems "rejuvenates"
the lower hierarchical structures and causes nearly unbounded evolution
of the biological world.
One must not forget that natural selection at work at higher hierarchical
levels is a manifestation of an important mechanism that assures the interaction
of biological systems with the environment and helps their survival. But
those problems first of all are in the realm of kinetics.
Additional experiments could refine the theoretical model and stimulate
an effort to obtain additional proof that the second law can be applied
in its classical definition (as formulated by R. Clausius and J.W. Gibbs)
to explain the origins of life on Earth or elsewhere in the Universe.
The findings of hierarchical thermodynamics, specifically supramolecular
thermodynamics of quasi-closed systems, and the published data about the
variation of the chemical composition of living organisms during ontogenesis
and phylogenesis confirm the thermodynamic tendency of biological evolution
and aging processes. According to the thermodynamic theory and experimental
data, the specific value of the Gibbs function of the formation of supramolecular
structures of organisms during ontogenesis and phylogenesis tends to a
minimum. The principle of the stability of chemical substances of the supramolecular
structures of tissues makes it possible to understand the causes of the
nearly unlimited evolution of the biological world.
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Additional information can be found on the Internet
Some notions and terms used in the theory are at the site (References 3,5,7,9)
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