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Thermodynamic and information entropy are considered as two forms of total entropic process in biosystems.
The origination of complexity cannot be compensated only by thermodynamic entropy.
When and where in the past the entropy has been produced that is a payment for biological organization at present?
The idea is discussed that the genetic information is an instrument of entropy disproportioning in time.
The Second Low realization today cannot be without taking into account the information entropy in past generations.
art ic l e i nf o
a b s t r a c t
Article history:
Received 14 February 2014
Received in revised form
30 May 2014
Accepted 12 June 2014
Available online 20 June 2014
The article attempts to overcome the well-known paradox of contradictions between the emerging
biological organization and entropy production in biological systems. It is assumed that quality,
speculative correlation between entropy and antientropy processes taking place both in the past and
today in the metabolic and genetic cellular systems may be perfectly authorized for adequate description
of the evolution of biological organization. So far as thermodynamic entropy itself cannot compensate
for the high degree of organization which exists in the cell, we discuss the mode of conjunction of
positive entropy events (mutations) in the genetic systems of the past generations and the formation of
organized structures of current cells. We argue that only the information which is generated in the
conditions of the information entropy production (mutations and other genome reorganization) in
genetic systems of the past generations provides the physical conjunction of entropy and antientropy
processes separated from each other in time generations. It is readily apparent from the requirements of
the Second law of thermodynamics.
& 2014 Elsevier Ltd. All rights reserved.
Keywords:
Information entropy
Generating of new information
Disproportionation of entropy
Unsteadiness in genetic systems
Compensation for antientropic processes
1. Introduction
Discussion about implementation of the Second law of thermodynamics in living systems has always been a collision of diverse
points of view, especially when discussing the role of information
processes in such fundamental biological phenomena as growth,
embryogenesis, ontogenesis and evolution. The fact that genetic
information is correlated with the entropy production was
expressed in the question: "what is the entropy (and energy)
payment for information" (Romanovskiy et al., 1984).
Blumenfeld (1977) wrote: "According to the thermodynamic
criteria any biological system is not more ordered than a piece of
rock the same weight". However, a few lines later he admitted
formalism of his approach: "Orderliness of living matter and the
information it contained have a sense".
Prigogine and his school developed thermodynamics of
open nonequilibrium systems and formulated idea about the
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formation (origination) of the new elements of biological organization and the process of retention of existing ones, i.e. selected
and forming a functioning biological system elements. At the same
time it is intended that this correlation is lawful only if there is a
direct causal physical link between these two opposite classes of
processes, even if the mapped events spaced in time and space.
Real basis for giving a comprehensive biological sense to such a
correlation is a well-established fact of conjugated positive
entropy of biochemical transformations of substances of food with
high content of free energy and thermodynamically impossible
without such conjugating processes of formation of various
macroergic equivalents, synthesis of complex organic molecules,
biopolymers etc.
We consider that continuous mutation process in genetic
systems during the evolution is the process of information entropy
production. The resulting unsteadiness in the genetic information
system is a necessary condition for the generation of new
information. Transfer of gene complexes to the next generation
that is supported by the selection and provides better information,
is that causal physical link between the positive entropy events in
the past generations and antientropy processes in their descendants. In fact, this speculative correlation between entropy and
antientropy processes is that attempt to overcome the paradox of
contradictions between the emerging biological organization and
the growth of entropy. The problem is complex and cannot be
described by a simple formula.
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inherent in the system potential which determines the spontaneous course of the process.
This logical impasse will be overcome, if we apply macromolecular or genetic level of biological systems, especially since all
the events of this level occur within the described system. The
causal connection between the events of these two levels is not
in doubt. Remaining within the limits of the natural science in
describing the phenomena and processes, we must, as before, in
the case of the description of intracellular physico-chemical
transformations, nd the force that determines the direction of
the spontaneously owing embryogenesis.
Without delving into details of embryogenesis we will try to
highlight this biological phenomenon from the point of view of
the requirements of the second law of thermodynamics. Positive
thermodynamic entropy balance in each cell of a multicellular
organism is not associated directly with the emerging and
growing organization, orderliness of this higher level in the
hierarchy of a multicellular organism. Obviously, without the
cells possessing of some properties which are necessary for the
formation of a multicellular organism, the embryo will not occur
and the result is just a larger number of cells, although each of
them has an obligatory positive entropy production balance.
Apparently, it is actually not the thermodynamic entropy produced in the cells of the developing embryo. It does not
determine the direction of the events of the considered hierarchy's level of living systems. The acquisition of certain properties that gives to dividing cells the ability to remain connected to
each other is apparently relevant to the implementation of the
new genetic information which originated in the evolving eukaryotic cellsthe predecessors of multicellular organisms. Its generating processed in genetic systems of these cells which possess
acceptable degrees of volatility and the possibility of mistakes
when replicating the nucleotide sequences of genes (Chernavsky,
2001). It was the rst and may be the most decisive step on the
path of the multicellular origination. Each of these countless
steps was an act of a new information birth.
The second, not less important event in the germ cells of
primary multicellular organisms was, obviously, origination in the
genomes of these cells of absolutely new, specic genes whose
activity modulated features of the hereditary cells phenotype in
such a way that in a growing embryo the cells with different
functional specicity began to originate. There is no doubt that
this became possible when the original genome of zygote was
enriched with a considerable set of genes new genes which did
not exist in free-living eukaryotic cells. The formation of a wide
spectrum of phenotypically different cellular elements in the
developing embryo was (and is) based on the same initial genome.
Eukaryotic cells have acquired an extremely useful evolutionary
innovation which allows them rational and sustainable use of the
resources of their actually large genome converting certain genes
in silence state or returning them in active one. We discussed it
more elaborately in the work (Mitrokhin, 2013).
The process of new genetic information generating is directly
conjugated with the changes of entropy (both thermodynamic and
information) but, however, it should be recognized that there is a
fundamental difference between the thermodynamic entropy and
the information one in probabilities of occurrence of elementary
events which make sense of each entropy type. The growth of the
thermodynamic entropy results in processes of biochemical transformations which are accompanied with the dispersion of part of
the free energy of the participants (presented in the system or
entering into it). The probability of elementary acts is very high
because the reactions of this type come with the release of energy.
Part of this energy is retained and will be then implemented for
the processes of thermodynamically improbable events, such as
the synthesis of universal macroergic compounds.
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performed conserving function; they provided memory and accuracy of the genome duplication in generations (Shcherbakov,
2005a, 2005b; Rice and Chippindale, 2001). The primary nucleotide sequence is strictly retained but larger elements of the
genome (genes, complexes genes, promoters and other elements)
are shufed. Impressive recent achievements of comparative
genomics prove this observation. Deep conservatism of nucleotide
sequences of genes contrasts with higher mobile gene composition of genomes and, to higher extent, evolution instability of
genome architecture, i.e. arrangement of genes in genome
(Koonin, 2009, 2012; Novichkov et al., 2009; Koonin and Wolf,
2010). It provides (especially in gametogenesis) with the origination of new, unexpected genetic combinations. This process goes
in parallel with the regular (evolutionarily more primitive),
ancient mutagenesis at the level of single nucleotide substitutions.
At the stage of selection there is no any creativity about which,
for example, Ayala, 2007. There is either "applause" or "stunned
silence". The creativity took place on the scene of life when
emerged in genetic information system new information realized
in an unexpected turn of events in the cell, in the modied
phenotype. If we see natural selection only as its nal stage, i.e.
elimination of nonviable forms, and at the same time we are aware
of the fact that this process does not require the functioning of any
mechanisms and is implemented on its own, then we lose sight of
the two rst and the most important stages of Darwin's natural
selection that he dened as the nature of the organism. In other
words, a creative role is played not by the last stage of the whole
selection process, but by its rst phase evolving information
systems.
4. Concluding remarks
We observe in a living cell several parallel conjugated processes
the driving force of which is accompanying these processes
entropy in two forms: thermodynamic entropy and information
one. The processes of metabolism providing the formation of
necessary substrates and macroergic equivalents proceed due to
spontaneous oxidation of food substances which possess the free
energy and are accompanied with the production of a considerable
degree of thermodynamic entropy. This type of entropy compensates for those conjugated antientropic processes (such as muscle
contraction, active transport of substances, the biosynthesis of all
polymers etc.) that cannot proceed spontaneously.
Basic events of the metabolic level are sequence of physicochemical transformations of small molecules and determine the
system's aspiration system for the most probable statethermodynamic equilibrium. However, it does not occur for the reason
that the system is open for matter and energy and contains an
internal source of informationgenetic system of cells which is a
macromolecular complex and requires the constant reproduction
of its structure for current information cell maintenance and for
transfer of information to the future generation.
Process of operations on genetic material includes two essentially different components. The elementary acts of polymerization
of the predecessor monomers which possess a considerable
reserve of free energy ow spontaneously and are accompanied
with the production of thermodynamic entropy. The resulting
molecular orderliness (i.e. the process of polymerization) is evidence of the decrease of the entropy balance member which
reects its production within the system. This rst component of
the genetic system elements reproduction provides the formation
of its physical and chemical structure.
The second aspectits informational content, its specic
sequence which provides the functional organization of the
system is inherited to the newly synthesized descendant
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