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1336 Norwegian Journal of development of the International Science No 17/2018
CHEMICAL AND BIOLOGICAL EVOLUTION: THE PRINCIPLE OF SUBSTANCE STABILITY IN
ACTION
Gladyshev G.
Doctor of chemical sciences, professor Principal scientist N. N. Semenov Institute of Chemical Physics Russian Academy of Sciences, Department of Design, Russian Academy of Arts,
Moscow
Abstract
The principle of substance stability is a driving force of evolution, facilitating a smooth transition of chemical evolution to biological evolution and determining thermodynamic selection of chemically energetic substances in the evolution, phylogenesis, and ontogenesis of living beings. Evolutionary selection of purines seems to play a specific role in formation of nucleic acids due to the action of the principle of substance stability. A special role of oxygen and environment in metabolism and catabolism in metabolic cycles is pointed out.
Keywords: life, the origin of life, biological evolution, Darwinism, hierarchical thermodynamics, the principle of substance stability, the second principle, Gibbs free energy, gerontology, aging.
Introduction
The goal of this work is, first, to demonstrate with simple examples that hierarchical thermodynamics based on the principle of substance stability is able to explain and predict directional variations in chemical and biological evolution and, second, to verify the assertion that Nature tends to the maximum structural stability at the molecular and supramolecular levels.
The principle of substance stability, or the feedback principle, was formulated by the author of this work in [1]. Later, considerations concerning physical verification of the principle were presented. The considerations were based on simple concepts [2-5]. It was stated that each atom, molecule, an isolated supramo-lecular structure (or structure of higher hierarchy) has a potentially limited capability of participating in interactions with other atoms, molecules, and supramolecu-lar or other structures. It was assumed that if particle i of hierarchy j (or subhierarchy j) spent much energy for binding to another particle of the same hierarchy j, there remains rather little energy in this particle i for binding to other particles of its hierarchy or particles of higher hierarchies (j+1). For instance, if a molecule (formed by strong chemical bonds) is relatively thermodynami-cally stable, it is unable to form rather stable structures that arise during formation of supramolecular structures (aggregates). Isolated supramolecular structures of the lower hierarchy j (consisting of a relatively small number of molecules) can participate in formation of supramolecular j+1 structures (consisting of a large number of particles) in accordance with their limited energy capabilities. It is established that the principle of substance stability acts at all hierarchical levels and sublevels of systems in all hierarchies of living matter [3-5]. The principle of substance stability plays a decisive role at particular stages of chemical evolution. It is the driving force of the origin of life, biological evolution, phylogenesis, and ontogenesis.
The principle can be formulated as follows:
«During the formation or self-assembly of the most thermodynamically stable structures at the highest
hierarchical level (j), e.g., the supramolecular level, Nature, in accordance with the second law, spontaneously uses predominantly the least thermodynamically stable structures available from a given local part of the biological system, belonging to a lower level, i.e., the molecular level (j-1), and incorporates these unstable structures into the next higher level, i.e. the supramolecular level (j) ».
Nature tends to maximum stability
The thermodynamic theory of the origin of life, biological evolution, and aging of living beings states that Nature tends to maximum stability at all hierarchical levels [8, 9].
Hierarchical thermodynamics allows only the directionality of this trend to be revealed because the composition of evolving systems continuously changes, and there are no data on absolute specific Gibbs free energies of formation of molecules and chemical structures. However, accurate comparative estimation of this stability is possible in some cases, e.g., for molecules with a different structure but an identical atomic composition. This can be used to reveal the optimum molecular structure used by Nature in evolutionary transformations of the chemical and su-pramolecular composition of organisms.
Special role of purines in chemical and biological evolution
The principle of substance stability allows important interesting conclusions to be drawn. A comparison of the Gibbs free energy for formation of adenine and guanine in physiological conditions with the similar values for their analogues with the identical chemical (atomic) composition makes it possible to state that purines are minimally chemically stable organic substances chosen by Nature to form highly stable supra-molecular structures favoring optimum metabolism.
To confirm the above statement, we compare Gibbs free energies of formation of the compounds in question.
Adenine nh2
Chemical Formula C5H5N5 Standard Gibbs Free Energy of Formation (A f G'°) = 125.61kcal/mol [6] Standard Gibbs Free Energy of Formation (A f G°) = 71.58 kcal/mole (S) [7]
2-Aminopurine
Chemical Formula C5H5N5 Standard Gibbs Free Energy of Formation (A f G'°) = 84.95 kcal/mol [6]
Here A f G'° is the Standard Gibbs Free Energy of atomic composition. Nature employs adenine as a frag-
Formation in physiological conditions, and A f G° (AGP298) is the Standard Gibbs Free Energy of Formation at 25 0C.
A comparison of the A f G'° values shows that adenine is far less stable in physiological conditions than 2-aminopurine though both compounds have an identical
ment of nucleic acids and ATP. It is hard to escape a conclusion that Nature uses the principle of substance stability and really tends to the maximum stability of supramolecular structures in evolution, phylogenesis, and ontogenesis.
A comparison of guanine and isoguanine yields a similar result.
Guanine
Chemical Formula C5H5N5O Standard Gibbs Free Energy of Formation (A f G'°) = 36.08 kcal/mol [6] Standard Gibbs Free Energy of Formation (A f G°) = 11.33 kcal/mole (S) [7]
Isogucmine
H
nh2
Chemical Formula C5H5N5O Standard Gibbs Free Energy of Formation (A f G'°) = 23.33 kcal/mol [6]
The comparison of guanine and isoguanine also (as the comparison of adenine and 2-aminopurine) shows that despite their identical composition, the above compounds are characterized by different stability. Guanine in physiological conditions has a considerably larger A f G'° than isoguanine. Consequently, guanine is chemically less stable than isoguanine. In this case, following the principle of substance stability, Nature also uses gua-nine as a comparatively unstable compound for formation of the stable supramolecular structure of nucleic acids.
A stability comparison of pyrimidine bases did not reveal the situation similar to purines. This allows a conclusion that purines as building blocks of life are of crucial importance for the processes of the origin and evolution of life. However, additional investigations are needed for verifying the conclusions drawn.
Oxygen favors development of life
Earlier the author formulated an approximate rule defining the trend of directional development of evolutionary processes resulting from enrichment of chemical compounds with oxygen [3, 4]. It was stated as follows: enrichment of substances with nitrogen atoms (with approximately unchanged ratio of other elements in the substance) often decreases their chemical stability, while enrichment of substances with oxygen atoms (with approximately unchanged ratio of other elements in the substance) increases their chemical stability. Though the "applicability scope" of the rule is rather vague, it allows comprehending some regularities of the directional evolutionary development. Indeed, metabolic cycles with the participation of oxygen tend to increase the amount of this element in tissues of living organisms, which ultimately leads to formation and excretion of stable compounds, such as CO2, H2O, urea, lactic acid, etc., from the organisms. This manifests the tendency of a living system toward maximum stability.
The above regularity represented as a directional tendency can be demonstrated by comparing the thermodynamic stability of Adenine - C5H5N5 and Guanine - C5H5N5O. The atomic compositions of these substances differ only by the presence of an oxygen atom in the guanine molecule. This difference leads to a substantial difference between the Standard Gibbs Free Energies of Formation (A f G'° and A f G°) of these substances: according to the approximate rule, guanine is more stable than adenine.
Another convincing example of the effect of oxygen on thermodynamic stability of substances is comparison of the ribose and deoxyribose structures. This example is considered in detail in a number of the author's investigations [4]. Replacement of -OH groups by carboxyl groups also leads to a higher stability of substances [7, 10]. This fact is often mentioned in study books.
The above rough rule needs commenting. The rule ignores the fact that various molecular groups containing nitrogen and oxygen make sometimes appreciably different contributions to the variation in the free energy of formation of metabolite molecules [10]. For example, in a few described cases contributions of molecular nitrogen-containing fragments (e.g., -NH3+ =NH2+, =N) are negative, which points to their relatively high stability. However, in most cases the above-mentioned contributions are positive, which agrees with the rule in its stating that nitrogen in molecules contributes to a decrease in stability of substances.
At the same time, contributions of oxygen-containing molecular fragments in the cases under consideration [10] are always negative, which confirms the tendency predicted by the rule that oxygen in molecules of metabolites increases stability of substances. It follows from this discussion that the above rule is effective within fairly narrow limits. For more rigorous estimation of A f G'° and A f G° one should be guided by the comprehensive study [10] and similar researches.
Thus, Nature tends to chemical stability of substances under evolution, enriching them with oxygen. However, this is opposed by the principle of substance stability: the tendency toward supramolecular stability counteracts this by enriching living systems with a low-stable (energetic) nitrogen-containing chemical substance. Each hierarchical level tends to maximum structural stability [11-12] but is prevented from reaching it by the action of the principle of substance stability. Living structures of all hierarchical levels are in states close to internal equilibrium but never reach equilibrium. This is an indication of life being manifested as a complex energy-dependent process close to equilibrium.
Appearance of oxygen in the Earth's atmosphere sharply activated evolutionary processes and changed the character of food of living beings. The rate of evolution increased under the effect of not only kinetic but also thermodynamic factors. In the general case, it seems to be promising to investigate evolutionary
transformations using the "chromatographic model" of evolution and aging, which studies the relationship of retention times and thermodynamic factors of sorption (absorption) of metabolites in metabolic processes. However, additional investigations are needed to discriminate the above-mentioned kinetic and thermodynamic factors. Nevertheless, examples are known that confirm that the low rate of metabolism caused by the low concentration of oxygen in the environment favors long life of organisms. These observations undoubtedly agree with hierarchical thermodynamics, which determines the slowed-down rate of metabolism. A classic example of the manifestation of this regularity can be the long life span of the naked mole rat (Heterocephalus glaber), which is known as a long-living mammal. Recall that thermodynamics also manifests its effect through food and all environmental conditions.
Recommendations that do not comply with thermodynamics
Science has become a mass phenomenon. Trying to make a discovery in life sciences a great deal of amateur researchers neglects world cognition methods that evolved for centuries. There is obvious underestimation of promise that physicochemical approaches, first of all thermodynamic methods, hold for explanation and prediction of phenomena using physical models of the origin of life, its evolution, and aging of living beings. Many colleagues believe that only attraction of investments will ensure success in researches aimed at expanding the life span. Indeed, much money is needed for empirical observations. But first and foremost the researchers need natural science education, which is often replaced by knowledge in the field of computer technologies.
In gerontology and food sciences there are recommendations that are based on the experience and individual views of a researcher and disagree with thermodynamics. It is often rather difficult to reveal these delusions because thermodynamics of aging science
usually identifies only trends in development and behavior of organisms. Nevertheless, there are recommendations of the thermodynamic theory based on strict quantitative estimations. It is always reasonable to take these recommendations into account.
From the position of the thermodynamic theory of aging, one can give a general recommendation: the search for geroprotectors, first of all, should begin with compounds - metabolites that contain structural groups characterized by a positive contribution to the Gibbs free energy of the formation of these substances, A f G'°
[10]. Such advice should be considered when using various food supplements and diets. However, one must bear in mind that the accuracy of the thermodynamic prediction depends on the capabilities of the A f G'° estimation model used.
Extended Darwin's selection
The thermodynamic theory of the origin of life, its evolution, and aging of living beings is a foundation for extended Darwinism. All transformations in the living world are associated with spontaneous processes governed by the second law of thermodynamics and nonspontaneous processes initiated by the environment
[11]. Selection in evolving chemical and biological systems proceeds under the directional effect of the principle of substance stability, which keeps dynamic systems in the state close to internal equilibrium. This manifests the tendency of all dynamic hierarchical quasi-equilibrium structures toward maximum internal stability [12], which they, strictly speaking, never achieve because of continuous substance and energy exchange in living systems.
Darwin's selection is undoubtedly directed by the thermodynamic principle of substance stability, which is physically verified at a quite acceptable level [4, 5].
Figure 1 shows the spiral of evolution, which is actually a physical and artistic symbol of its thermodynamic directionality.
Credit: U.S. Geological Survey Department of the Interior/USGS U.S. Geological Survey/photo by Jane Doe
Fig. 1- Spiral of evolution, the symbol of its thermodynamic directionality
Conclusions
Estimations of stability of chemical and biological structures in the course of chemical and biological evolution confirm the effective directive force of the principle of substance stability. This principle determines the composition and structure of nucleic acids and explains why Nature seems to prefer selection of purine bases and ATP as thermodynamically significant structures of the biological world. Thermodynamics explains why appearance of oxygen in the Earth's atmosphere contributed to the burst-like development of life on the planet. The thermodynamic approach helps reveal probable factors governing the life span of organisms.
The author hopes that this material is easy to comprehend for any well-educated physical chemist, biochemist, organic chemist, and common specialist acquainted with chemical and biological thermodynamics.
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DYNAMICS OFSETTLING OF THEVYATKARIVER BASIN BYWILD BOAR IN
Dvornikov M.
Doctor of biological sciences, leading researcher Professor Zhitkov Federal State Budgetary Russian Research Institute of Game Management and Fur
Farming, Kirov
ДИНАМИКА ЗАСЕЛЕНИЯ КАБАНАМИ (SUS SCROFA L.) БАССЕЙНА РЕКИ ВЯТКА
Дворников М.Г.
д.б.н., ведущий научный сотрудник Всероссийский научно-исследовательский институт охотничьего хозяйства и звероводства им.
проф. Б.М. Житкова, Россия, г. Киров
Abstract
Due to changes in climate and in plant cover the dynamics of settling of the Vyatka River basin by wild boar (Sus scrofa L.) was studied. The data on the content of the bone material in sediments on the sites of hunters' stays in Mesolithic-Neolithic-Bronze and Iron epochs are given. Current materials on the ecology of wild boar show that its vital functions in natural ecosystems always depend on oak (Guercus robur L.) distribution. Anthropogenic transformation of landscapes favours its survival in taiga ecosystems. With the growing bioclimatic trend and moving of borders of broad-leaved forests to the north the increase of wild boar numbers is expected.
Аннотация
В связи с изменениями климата и растительного покрова, рассмотрена динамика заселения кабанами (Sus scrofa L.) бассейна реки Вятка. Приводятся сведения по содержанию костного материала в отложениях на стоянках охотников эпохи мезолита-неолита-бронзы и железа. Обобщение современных материалов экологии кабанов указывает на то, что его жизнедеятельность в природных экосистемах всегда зависела от распространения дуба (Guercus robur L.). Антропогенные преобразования таёжных ландшафтов способствуют его выживаемости. Предполагается увеличение поголовья кабанов с нарастающим биоклиматическим трендом и сдвигом границ широколиственных лесов на север.
Keyword: climate change, wild boar, Vyatka river basin, transformation of taiga landscapes.
Ключевые слова: изменение климата, кабан, бассейн реки Вятка, преобразования таёжных ландшафтов.
