Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis)
An unconventional approach to the study and analysis of ontogenesis, based on the duality of its nature is proposed: in the first case this is a phenomenon, in second case it is a process. As a phenomenon, ontogenesis covers all manifestations of living, providing the continuity of life on all the l...
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irk-123456789-1097432016-12-13T03:02:11Z Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) Kovtun, M.F. An unconventional approach to the study and analysis of ontogenesis, based on the duality of its nature is proposed: in the first case this is a phenomenon, in second case it is a process. As a phenomenon, ontogenesis covers all manifestations of living, providing the continuity of life on all the levels of its organization, ensures lifespan of a species, regardless of the individual‘s life, and thus the dynamic stability of the cycle of matter and energy flows, the evolution cycles, the biosphere and life in general. As a process it is the implementation of ontogenesis by the way of successive morphogeneses. There is a gap between these components of ontogenesis (the phenomenon and the process) that must be filled out. This is a complex task in the scope of the ontogenesis evolution problem (making the theory of ontogenesis). Предложен нетрадиционный подход к изучению и анализу онтогенеза, исходя из двойственности его природы: с одной стороны, это явление, с другой — процесс. Как явление онтогенез охватывает всю живую природу, обеспечивая непрерывность жизни на всех уровнях её организации; обеспечивает продолжительность жизни вида независимо от жизни индивида и тем самым динамическую стабильность круговорота вещества и потоков энергии, эволюции круговоротов, биосферы и жизни в целом. Процесс — реализация онтогенеза путём последовательных морфогенезов. Между этими составляющими онтогенеза (явление и процесс) — свободное пространство; заполнить его и представляет собой комплекс задач в рамках проблемы эволюции онтогенеза (создание теории онтогенеза). 2013 Article Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) / M.F. Kovtun // Вестник зоологии. — 2013. — Т. 47, № 3. — С. 195-204. — Бібліогр.: 22 назв. — англ. 0084-5604 DOI 10.2478/vzoo-2013-0019 http://dspace.nbuv.gov.ua/handle/123456789/109743 591.3 en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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An unconventional approach to the study and analysis of ontogenesis, based on the duality of its nature is proposed: in the first case this is a phenomenon, in second case it is a process. As a phenomenon, ontogenesis covers all manifestations of living, providing the continuity of life on all the levels of its organization, ensures lifespan of a species, regardless of the individual‘s life, and thus the dynamic stability of the cycle of matter and energy flows, the evolution cycles, the biosphere and life in general. As a process it is the implementation of ontogenesis by the way of successive morphogeneses. There is a gap between these components of ontogenesis (the phenomenon and the process) that must be filled out. This is a complex task in the scope of the ontogenesis evolution problem (making the theory of ontogenesis). |
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Kovtun, M.F. |
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Kovtun, M.F. Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) Вестник зоологии |
author_facet |
Kovtun, M.F. |
author_sort |
Kovtun, M.F. |
title |
Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) |
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Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) |
title_full |
Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) |
title_fullStr |
Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) |
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Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) |
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ontogenesis: a phenomenon and a process (on the problem of the evolution of ontogenesis) |
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Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2013 |
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http://dspace.nbuv.gov.ua/handle/123456789/109743 |
citation_txt |
Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis) / M.F. Kovtun // Вестник зоологии. — 2013. — Т. 47, № 3. — С. 195-204. — Бібліогр.: 22 назв. — англ. |
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Вестник зоологии |
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AT kovtunmf ontogenesisaphenomenonandaprocessontheproblemoftheevolutionofontogenesis |
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2025-07-07T23:35:57Z |
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1837033175859068928 |
fulltext |
UDC 591.3
ONTOGENESIS: A PHENOMENON AND A PROCESS
(ON THE PROBLEM OF THE EVOLUTION OF ONTOGENESIS)
M. F. Kovtun
Schmalhausen Institute of Zoology, NAS of Ukraine,
vul. B. Khmelnytskogo, 15, Kyiv, 01601 Ukraine
E-mail: kovtun@izan.kiev.ua
Ontogenesis: a Phenomenon and a Process (on the Problem of the Evolution of Ontogenesis). Kovtun M. F. –
An unconventional approach to the study and analysis of ontogenesis, based on the duality of its nature
is proposed: in the first case this is a phenomenon, in second case it is a process. As a phenomenon, onto-
genesis covers all manifestations of living, providing the continuity of life on all the levels of its organi-
zation, ensures lifespan of a species, regardless of the individual‘s life, and thus the dynamic stability of
the cycle of matter and energy flows, the evolution cycles, the biosphere and life in general. As a pro-
cess it is the implementation of ontogenesis by the way of successive morphogeneses. There is a gap between
these components of ontogenesis (the phenomenon and the process) that must be filled out. This is a
complex task in the scope of the ontogenesis evolution problem (making the theory of ontogenesis).
Key wo rd s: ontogenesis, phenomenon, process, morphogenesis, cycle of matter and energy, biologiza-
tion, individualization, essence.
Îíòîãåíåç: ÿâëåíèå è ïðîöåññ (ê ïðîáëåìå ýâîëþöèè îíòîãåíåçà). Êîâòóí Ì. Ô. – Ïðåäëîæåí íåòðà-
äèöèîííûé ïîäõîä ê èçó÷åíèþ è àíàëèçó îíòîãåíåçà, èñõîäÿ èç äâîéñòâåííîñòè åãî ïðèðîäû: ñ
îäíîé ñòîðîíû, ýòî ÿâëåíèå, ñ äðóãîé – ïðîöåññ. Êàê ÿâëåíèå îíòîãåíåç îõâàòûâàåò âñþ æèâóþ
ïðèðîäó, îáåñïå÷èâàÿ íåïðåðûâíîñòü æèçíè íà âñåõ óðîâíÿõ å¸ îðãàíèçàöèè; îáåñïå÷èâàåò ïðî-
äîëæèòåëüíîñòü æèçíè âèäà íåçàâèñèìî îò æèçíè èíäèâèäà è òåì ñàìûì äèíàìè÷åñêóþ ñòàáèëü-
íîñòü êðóãîâîðîòà âåùåñòâà è ïîòîêîâ ýíåðãèè, ýâîëþöèè êðóãîâîðîòîâ, áèîñôåðû è æèçíè â
öåëîì. Ïðîöåññ – ðåàëèçàöèÿ îíòîãåíåçà ïóò¸ì ïîñëåäîâàòåëüíûõ ìîðôîãåíåçîâ. Ìåæäó ýòèìè
ñîñòàâëÿþùèìè îíòîãåíåçà (ÿâëåíèå è ïðîöåññ) – ñâîáîäíîå ïðîñòðàíñòâî; çàïîëíèòü åãî è ïðåä-
ñòàâëÿåò ñîáîé êîìïëåêñ çàäà÷ â ðàìêàõ ïðîáëåìû ýâîëþöèè îíòîãåíåçà (ñîçäàíèå òåîðèè îíòî-
ãåíåçà).
Êëþ÷åâûå ñ ëîâ à: îíòîãåíåç, ÿâëåíèå, ïðîöåññ, ìîðôîãåíåçû, êðóãîâîðîò âåùåñòâà è ýíåð-
ãèè, áèîëîãèçàöèÿ, èíäèâèäóàëèçàöèÿ, ñóùíîñòü.
Introduction
The ontogenesis or individual development is the process that covers the time from the beginning to the
end of the life cycle of individuals (organisms). It is divided into prenatal period or embryogenesis (prior to
withdrawal from embryonic membranes) and postnatal period of development, or post embryogenesis (after
leaving the embryonic membranes). The embryogenesis is an intimate process hidden from the direct obser-
vation and its study requires special methods. The studies of embryogenesis have a long history; we can find
fairly complete information about the chick embryogenesis already in the works of Aristotle (384—322 BC).
Later it became apparent that the embryological studies do not provide information about the ontogen-
esis in general and that studying processes of individual development requires knowledge from other fields such
as genetics, cytology, molecular biology, biochemistry, etc. With the involvement of these sciences, and devel-
opment of the experimental trend in embryology, the classical embryology relays race to a new branch, the
biology of individual development, with a wider range of tasks, including the origins and patterns of ontoge-
nesis evolution. Until now, there is a question in focus, if the unicellular dividing can it be considered to be
the ontogenesis. A. N. Severtsov (1939) denied the existence of the ontogenesis for the unicellulars, calling it
the histogenesis, and A. K. Dondua (2005) was close to Severtsov’s attitudes. However, most biologists study-
ing ontogenesis expressed the opposite opinions (Smith, 1968; Dogiel, 1951; Korotkov, 1979, 1991; Desnitsky,
2006; Dovgal, 2010). In particular, I. V. Dovgal believes that “…the mechanisms of ontogenesis regulation at
the cellular level in the protozoans could be a model not only for the formation of the multi-cellular mecha-
nisms of morphogenesis (as in Volvox), but for the mechanisms, which are similar to those occurring in the
fertilized egg in prior to cell division.” G. P. Korotkova (1979) presented a selection of data, showing in her
Vestnik zoologii, 47(3): e-1—e-10z, 2013
DOI 10.2478/vzoo-2013-0019
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opinion the origin of the processes that lead to reproduction and ontogenesis on the stage of chemical evolu-
tion. Of course, ontogenies of the protozoans, multicellular invertebrates and vertebrates are different, but their
origins are obviously common.
Ontogenesis of all living organisms is a product of biological reproduction and morphogenesis. The fer-
tilization and biological reproduction are starting ontogenesis (launch the process), and the morphogenesis is
a tool to implement it. As already mentioned the ontogenesis cannot be understood only on the basis of embryo-
genesis or postembryogenesis studies.
However, we can not ignore the fact that the study of embryogenesis is almost the only source of empir-
ical data, which shows the development of the organism before it leaves the embryonic membranes. Due to
specific embryogenesis studies, we have got information about their variability (intra-and inter-specific), about
the differences in the process behavior in phylogenetically closely and distantly related species, the heterochronies,
heterotopias, embrionization, pedomorphosis, equifinalism, etc. Therefore, the specific embryogenesis studies
still remain in demand in the XXI century, however, with the reconsidering (upgrading) of the purpose and
objectives of research. The embryology, as the part of the biology of individual development, should have a
general biological (theoretical) problem statement: transfering to (specific) ontogeneses origin and evolution,
variability and stability through the study of specific embryogeneses, i. e., the solution of the tasks that form
the basis of the theory of ontogenesis.
In this paper, we have the groundwork’s of specific embryogenesis. Relying on literature data, we propose
an approach to the discussion of the problem of ontogenesis, proceeding from its duality. In particular, we believe
that in ontogenesis must distinguish at least two components: ontogenesis as a phenomenon and as a process.
Ontogenesis as a phenomenon and as a process
We consider ontogenesis as a phenomenon in the wide sense, akin to gravity, mag-
netism, radioactive emission or, finally, life, and taking into account its significant dif-
ferences. If the gravity, magnetism, radiation levels, subject to the physical laws of mat-
ter are inherent in general, in case of the ontogenesis it is inherent in the living matter
and, only starting from a certain level of its organization. Due to that the ontogenesis as
the phenomenon covers all the animate nature, providing the continuity of the life of organ-
isms at all levels of the organization. Process is the ontogenesis implementation by suc-
cessive morphogeneses.
The ontogenesis as the process is available to study with the different methods of var-
ious sciences, including the experimental approach (the leading role remains for the embry-
ology), the ontogenesis as the phenomenon remained out of research scope: there are no
direct methods for investigation of this manifestation. It is possible to analyze, discuss,
debate, based on the general scientific methods (techniques) as the extrapolation, anal-
ogy, philosophy…
The phenomenon is a philosophical category, and its dialectical pair is the essence.
It is believed that the knowledge proceeds from the phenomenon (simplified – the detec-
tion of an object or thing) to the disclosure of (knowledge) its essence (internal connec-
tions and external characteristics consist the essence of the subject (Soviet…, 1984).
It is not correct to talk about the discovery of ontogenesis as a subject (if not to con-
sider the naming of this phenomenon as “ontogenesis”): it exhibits itself in the “race relay”
from generation to generation of organisms and maintaining the essential qualities of the
species and has done so for millions of years and thousands of generations.
The phenomenons are not single-valued, both in origin and in time they may be “eter-
nal”, and also can be created by the nature and mind. For example, rain and lightning,
tides, gravity, cosmic radiation are the “eternal” existing phenomenons. Cognition of the
essence of these phenomenons has led to the creation of artificial sources of energy and
devices that convert energy from one form to another, obtaining the materials with desired
properties, allowed humanity to overcome the force of gravity, to get into space, to the
depths of the ocean, etc.
The ontogenesis refers to the phenomenons created by nature, i. e., evolution. The
time of its “creation” in general terms, obviously, coincides with the beginnings of life,
and more specifically – with the beginnings of individuality.
There is a gap between these two components of ontogenesis (the phenomenon and
the process) and, obviously, the filling this gap is a set of tasks in the scopes of the onto-
genesis problem (a theory of ontogenesis).
e-2 M. F. Kovtun
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Ontogenesis as a phenomenon, regardless of the length of life of the individual organ-
ism, manifests in a longer or shorter lifespan of an individual, maintaining the essential char-
acteristics of organisms that consist this species (taxon) including the presence of variation;
and diversity in a variety of the animate nature (biodiversity), and in the strong stand of
the whole life on the Earth. Ontogenesis as the process manifests in the prenatal ontoge-
nesis and its variability, time of sexual products maturation, variability of timing of organs
maturation in relation to the demand for their roles in the postnatal period, types of divi-
sion, cyto- and histogenesis; life of the individual organism and the embryonization.
Lifespan is obviously controlled by the cycles of matter and energy flows, and
depends on the role of the species (life form) in the cycle. Therefore biosphere preserves
all types of organisms with different lifespans and various functions to maintain the dynam-
ic of the cycle. Some of them, with a short life cycle, are the operative components of
the cycle, while others, with long lifespans, are reservoirs or storage of excessive energy
and matter (Kovtun, 2006).
During the evolution of ontogenesis the mechanisms of its changes or modifications
(restructuring) were developed. The main known changes are the changes of embryoge-
nesis length and interrelations of the embryogenesis periods; rescheduling the anlage, for-
mation and maturation of organs (heterochrony), change in the topography of anlages
(heterotopia), the development of embryonic membranes, improve embryo protection and
trophics; transition to the indirect ontogenesis and vice versa; acceleration of maturing
of gametes (neoteny), internal fertilization, and also addition stages to the ontogenesis
(anaboly) and care of offspring. Of course, some of these “mechanisms” influenced the
ontogenesis as the phenomenon, but mostly, the ontogenesis as the process was recon-
structed (i. e., morphogeneses).
Originally the ontogenesis was being formed as directed process, and his ultimate
goal – to ensure the reproduction of the offsprings “of its own kind”, which could per-
form the functions in the biosphere previously performed by its ancestors.
Returning to the dual nature of the ontogenesis (as a phenomenon and as the pro-
cess) and the philosophical interpretation of the categories of “phenomenon” and “enti-
ty” we should focus our attention on the well-known definition: penetration into the phe-
nomenon is possible only through studying it. There is a mandatory logical connective
between the categories phenomenon and entity – learning process (fig. 1).
As can be seen in the scheme, the process of cognition of the ontogenesis is multi-
ple-vector. Paleontology gives useful information for the ontogenesis analysis (recapitu-
lations) (Shishkin, 2012). The science anticipates a genius, who will be able not only to
summarize the knowledge of all the sciences, but also see something in the phenomenon,
that is no one have seen before. Is it possible? Let us leave it unanswered. It is to recall
the statement of G. Yu. Lubarsky on the situation in science, which is emerging because
of the unfocused facts accumulation: accumulation of the facts is excused because one
day the theorist would come and sort it in order, make generalization and a theory.
“However, even if such a theorist will appear, he will be gasping for the lack of evidences
e-3Ontogenesis: a Phenomenon and a Process...
Fig. 1. Process of ontogenesis cognition.
Ðèñ. 1. Ñõåìà ïðîöåññà ïîçíàíèÿ îíòîãåíåçà.
The process
of cognition
The phenomenon
(of ontogenesis)
The essence
(of ontogenesis)
philosophy
embriology
biochemistry cytology
genetics
physics
chemistry
Biology of individual development
mol. biology
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“in purely empirical articles” (Lubarsky, 1994). This suggests there is a methodology defi-
ciency in the ontogenesis study. However, each of the sciences has certain developments
in the study of this issue.
Geochemical and biochemical aspects of the origin of ontogenesis
It will be logical to assume that the nascent living matter could exist and evolve, in
the case if it could reproduce itself. Therefore, the mechanisms of reproduction evolved
in parallel with the evolution of life, which is reflected in the famous thesis “a living from
the living”. The question is: where did the life come?
There are two alternative views on the origin of life on Earth: it was conceived and
created by the Creator or it emerged naturally as a result of geochemical, biochemical
and organic evolution of matter. The most coherent theory of the life origin is Oparin
and Haldane’s theory, which has both supporters and opponents. On this subject, there
is an extensive literature that indicates unresolved problems.
The monograph by G. P. Korotkova (1979) contains enough proper data review and
analysis of the “chemical” origin of life and ontogenesis, assertions on this issue the author
argues on the basis of the works by V. Vernadsky, A. Oparin, M. Kamshilova, D. Bernal
and others.
Obviously, all judgments about the origin of life and ontogenesis are more or less well-
reasoned hypotheses, regardless of confirmation or refutation by the experimental data.
So rightly is to suggest other hypotheses that arise in the discussion of the existing ones.
There is no credible evidence for that the heavenly bodies or planetary systems can
produce “their own kind”. However, the assumption of a plurality of, for example, solar
systems could create assumption, of their reproduction. Inexhaustible number of chemical
elements in the Earth’s history (C, H, Fe...) suggests their reproduction in the period of
history, which was before the occurrence of organics. Also the possibility of origin of the
new elements from the existing is known, for example, the formation of the helium from
the combustion of hydrogen. Hence the reproduction is the immanent (and permanent),
property of Nature, and all that occurred in nature, endowed with the ability to reproduce.
This “power” has been changed and intensified (evaluated) in the transition from the organ-
ic geochemical evolution, elaborated and improved the mechanisms of reproduction.
Consider another aspect. In the geological epoch of Earth’s history solar energy falling
on the Earth, was “wasted” (chemical reactions seems to occur by the internal energy
of the Earth). Cycling of matter and energy flows (geochemical period), apparently were
still in its infancy and had a local character. Earth was “interested” in the appearance
of consumers and transmitters of this energy. That situation stimulated transition from
the geochemical to biochemical and organic evolution. The growing cycle of matter and
energy have also was “interested”. Synthesized organic matter, and later probionts,
bionts, coenoses, primitive organisms, which consumed energy from the Sun, not only
converted it into energy of life, but over time started to accumulate that energy.
They stepped up the metabolism, keeping the solar energy at the surface of the Earth
and in the waters of the ocean, bringing the significant changes in the circulation of mat-
ter and energy, raising it to the level of the biosphere to coenotic, and accelerating evo-
lutionary processes. As V. I. Vernadsky (1930, 1967), and later M. M. Kamshilov (1970)
suggested, the components of the cycle, becoming biological, became individuals.
Individuals became real bearers of ontogenesis.
A few remarks to the problem. The energy, as well the matter, are carrying the infor-
mation. (To some extent this assertion does not match the cybernetic interpretation of
the information.) Hence: the information is also included in the cycling of matter and
energy flows, and in it, as in the cycle, also contains information on how the transfor-
mation (evolution) of matter, energy and circulation in case of a particular planet. We
can not agree with the authors (e. g. Goodwing, Lennox, 2007), who believe that the infor-
mation carrier is only the mind. Are bricks which are lying on the road, or a boulder in
e-4 M. F. Kovtun
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the forest, do not have the information? Then on which basis of paleontologists, and foren-
sic evaluators draw their own conclusions?
Mind retrieves the information, analyzes it, converts it, and creates new informa-
tion on this basis. That is, the mind, from our point of view, is the secondary creator and
bearer of information, when the first is the matter and energy.
The interesting idea about the properties of the information was expressed by
A. A. Lyapunov (1965). In particular, the information is not subject to the law of con-
servation. The information, as opposed to matter and energy can be re-created or irre-
trievably lost (cited by Eskov, 2000). “Lost information”, apparently, is the information
about the technology of construction of the pyramids, and... of the origin of life. The task
of the mind is not only to create a new information, but also to reconstruct the lost knowl-
edge, and the science is based on it.
In this section we intended to show that it is unproductive to deny the possibility
of the initiation of the processes leading to the evolution of life on the Earth for the geo-
chemical stage. Indeed, a combination of chemical elements that exist on the Earth orig-
inally led to materials with any desired properties. These materials have been created by
human (by human mind) who have studied the essential properties of the original mate-
rials in the last 100—200 years. So why do the chemical reactions and recombination of
the original elements with the third forces (catalysts, autocatalysis, heterocycles, solar ener-
gy) could not lead to organic, self-organizing systems, and later to the life, having on it
indefinitely time (billions of years), energy and materials? V. N. Snytnikov (2006) sub-
stantiates a very bold idea of the “astrocatalysis”, which states that “...the primary
organic compounds were synthesized in the protoplanetary circumstellar environment,
as planets form in areas of the catalytic synthesis of organic compounds”.
Role of the cycle of matter and energy in the evolution of ontogenesis
V. I. Vernadsky believed that life was not orignally existing in the form of organ-
isms (individuality), but as a matter of the biosphere.
Among the trends in the evolution of biosphere V. I. Vernadsky listed the accumu-
lation of energy, emergence of new forms of the chemical elements migration, tenden-
cy of biogenic migration of atoms to a maximum, increasing role of living matter in for-
mation and stabilization of the Earth surface.
V. I. Vernadsky assigned the crucial role in the current system in the biosphere to
the cycling of matter and energy. That means that the cycle, escaping from the entropy,
is in constant need of intensifying the biogenic migration of atoms, which, in turn, requires
new forms of migration and expansion of the arsenal used in the biosphere resources.
In our view, the “new forms of migration of atoms” can be achieved mainly by the
life forms, and only they are able to intensify the biogenic migration of atoms (Kovtun,
2006). Naturally, the dynamic stability of the cycle could be maintained by the stable
“work” of its components. Such stability is inherent for the components that can play
its unique role in the cycle and in the whole biosphere for more or less long time, per-
fecting it in a changing environment (under the control of natural selection and the cycle
of matter and energy). The “components” (biont, coenobionts, organisms) preserve and
maintain the required level of their unique function only by the reproduction of “their
own kind” and giving them their functions in the cycle. The mechanism of transmission
is the secret under seven seals. However, there are traces that leads from the period of
chemical evolution, the chemical cycle of matter and energy (Korotkov, 1979).
Life as the biosphere (Vernadsky, 1967), coenoses (Korotkova, 1991) or ecosystems
(Eskov, 2000) had an abstract character with a high degree of the heteromorphic ele-
ments, composing coenoses. Heteromorphicity implied the heterofunctionality of certain
elements, and this is a “material” for natural selection. The question is, how the “pre-
cursor elements” for keeping the system cycles, were able to transfer their (possibly unique)
e-5Ontogenesis: a Phenomenon and a Process...
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functions in coenoses to the “child elements”? Or you must admit, that “immortality”
of the elements that based on their “chemical” nature should not be taken absolutely unre-
al. If you recognize the latter, we should admit the fact that ontogenesis, as such, to the
coenosis stage of evolution of life, has not been formed yet. (There are statements about
“immortality” of akaryote unicellulars; Eskov, 2000.)
In the works of these authors, there is the thesis that the individualization of life comes
through the biologization of cycle components. The inevitable consequence of the
acquirement of biological function was the loss of “immortality”. The very urgent task
is a transmission to the biological components the cycles of their (unique) features and
functions in coenosis, biosphere to the next generation “components“. During this peri-
od, a phenomenon called “ontogenesis” appeared. As the ontogenesis arose in the pro-
cess of “biologization” of “non-biological” components of the cycle, it was bound to use
the mechanisms, which existed among the non-biological components of coenoses.
Others did not exist.
Let us return to the heteromorphic and heterofunctional coenoses, that offered great
opportunities for natural selection. It is natural to think that the natural selection was pro-
ductively (constructively) worked in the “original chaos”, as it had some clear guidance
or tasks. These “guidelines” were defined by the needs of the biotic cycle of matter and
energy. That means the natural selection had to work “for hire” in the cycle, selecting
bionts and then individuals with the necessary at this time functional, biochemical prop-
erties, capable of increasing biogenic migration of atoms and the emergence of “new forms
of migration of chemical elements” (Kovtun, 2006).
While selecting the functional properties of the individuals, selection was also simulta-
neously (indirectly) found the transmission mechanisms of these properties to the offspring.
Therefore, the individuals has recieved the mechanism of offspring properties transferring,
as a set of mechanisms inherent the coenoses elements, components and cycles (i. e., orig-
inally having chemical nature). A mechanism of unique properties transmission to offsprings
and mechanisms of reproduction were improved through the trial-and-error process.
Obviously it can be argued that ontogenesis as a phenomenon was formed in the biol-
ogization of circulation components up to individualization of bionts. Further evolution
occurred by improving morphogeneses those implemented the ontogenesis, i. e., the com-
ponent of the ontogenesis, which we call the process of its implementation.
Thus, we believe that there is reason to consider the cycles of matter and energy
(chemical, biochemical, biotic, or a “system of cycles”, as V. I. Vernadsky called them)
initially “are interested” to form individuality (and consequently, the reproduction
mechanisms) as stable and reliable components to ensure the continued functioning of
e-6 M. F. Kovtun
Fig. 2. Factors that initiate ontogenesis germination ( ) and ontogenesis ( ) derivatives.
Ðèñ. 2. Ôàêòîðû, èíèöèèðóþùèå çàðîæäåíèå îíòîãåíåçà ( ), è ïðîèçâîäíûå îíòîãåíåçà ( ).
ONTOGENESIS
Evolution
(geochemical, biochemical, organic)
Reproduction
Cycle of matter and energy
Age, cycle demand for
intensification of biogenic
atom migration
Cycle dynamics
and life stability
Biologization of
cycle componentsContinuity of life
(evolution)
Individualization
Independence of
species lifetime
regardless of
individual lifetime
Natural selection
Biodiversity
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the cycle and the whole biosphere (escape from entropy). If so, then the cycle of mat-
ter and energy flows is one of the leading creators of the ontogenesis (as a “phe-
nomenon” and “process”). Natural selection was the tool of the cycle (fig. 2).
Variability and stability of ontogenesis
Duality of the ontogenesis (as a phenomenon and process) is shown in its other char-
acteristics, in particular, in its variability and stability. M. A. Shishkin considers the stabil-
ity phenomenon of ontogenesis is an important characteristic: “...any concept, which could
be a new stage in the development of Darwinism, should be evaluated primarily by how it
is able to explain the origin of the stability of the normal ontogenesis” (Shishkin, 1988).
It seems that the variability of normal ontogenesis in its formation and evolution is
not less interesting, these phenomenons were parallel. (Variability as an intrinsic prop-
erty of the nature existed on the protobiological stage of evolution, so in the process of
“biologization” the components of cycle (bionts) “inherited” this variability.)
It only seems that nature of variability has less riddles. The origins of variability, prob-
ably lie in heteromorphic cenoses, protobionts, primary bionts; variety of physical and
chemical organization of the different parts of the Earth surface and in a well-developed
hybridization ability of early bionts (Korotkova, 1979); “ heterogeneous living matter always
observed in the biosphere, and life is always performed various biogeochemical functions
simultaneously“ (Vernadsky, 1930).
While analyzing the variability of ontogenesis, we cannot ignore the variability and
diversity of its components (the mechanisms giving a start to ontogenesis and accompa-
nying it all over) as follows: the type of reproduction, fertilization, types of zygote
cleavage, gastrulation, types of cell cleavage, speed and timing of the prenatal ontogen-
esis flow, development (direct, indirect, etc.) In addition, each of these components has
its own nuances, and is variable in its turn. For example, the length of prenatal ontoge-
nesis in vertebrates is from a few days up to six years or more, the number of offspring
in a brood is from one to several tens of thousand, the number of broods per life is from
one (for salmonid fish) to ten or more, the number of broods per season is from zero to
3—4; procreation is the viviparity, oviparity, oviviparity etc.
Equally diverse is the asexual reproduction: the binary division in the bacteria,
unicelullar algae, fungi, the architomy and paratomy of the worms, the formation of
zoospores in multicellular algae, spores in mosses and ferns; internal kidneys of some mul-
ticellular invertebrates, etc. Possibility of transition from asexual to sexual reproduction
and vice versa is also important, as well as parateny, hermaphroditismy, monosexuality:
males are unknown in one of the groups of echinoderms (Desnitsky, 2005). An exam-
ple demonstrating the duality of variability and stability of the ontogenesis is given by
P. G. Svetlov (1972): the developments of the ascidians by the sexual and asexual repro-
duction are profoundly different, but it is really difficult to distinguish the adult ascidia,
which was developed from the zygote by the embryogenesis, and the ascidia, developed
by budding, as they are identical.
The components discussed above, were acquired during the evolution of ontogene-
sis, but their variability refers to the evolution of ontogenesis as a process, i. e., morpho-
geneses, which realized ontogenesis as a phenomenon.
The original variability on the stage of protobionts and primitive bionts was very prob-
ably disordered and chaotic. The gradual evolution of life towards the individualization
eliminated “non-constructive” manifestations of variability. The biotic cycle and natu-
ral selection were becoming regulation factors. Since the selection can operate effective-
ly in the presence of environmental factors variation and heterogeneity of the coenoses
components (biosphere), then it maintains a “constructive” part of the variability and
scoops the carriers of function, able to intensify the “biogenic migration of atoms” and
expand the arsenal of used resources of the biosphere. In addition, as already was men-
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tioned, the natural selection as a tool of the cycle is likely not so much interested in a
form of organisms, but in their function in the biocoenosis and the biosphere, thereby,
the ability to do their “job” based on the needs of the cycle and the biosphere in a given
space and time.
The unique functions in coenoses, biosphere and in cycle can be performed only by
fully formed organisms (definitive), but not their embryos.
We suggest that these are the reasons of the morphogenetic variability of ontogen-
esis as a process (especially in its prenatal period) with the stability of the end function
and form of the reproduced product connected with it.
In this case the natural selection performs the search function, based on the needs
of the cycle, and the stabilization function for the form and function of the organism.
The evolution of ontogenesis (discussion)
This problem has a lot of questions and only a few answers of a hypothetical kind.
is the only axiomatic issues are that the living matter was able to reproduce itself from
the beginning, and that the ontogenesis, as such, should be formed with the development
of individuality...
If we follow the theory of the origin of life in a natural way, i. e., by evolution, then
we recognize the emergence of ontogenesis during this evolution, that is in the process
of biological function of matter.
Obviously, the most difficult part of the problem of ontogenesis are the conditions
and stimulating emergence of ontogenesis as a phenomenon and initiation of mechanisms
for its implementation.
Heredity is only a part of ontogenesis, one of the many. The mechanism of trans-
mission of hereditary properties has been discovered, but the emergence of this mecha-
nism still a mystery. “Complexity” of this issue makes us talking of the Creator or the
Mind, the only feasible solution of this problem. We believe that the extreme “complex-
ity” of the issue was dictated by the failure of our mind to create a self-reproducing cell,
which is functioning as a factory for production of something. However, the difficulty
disappears if we accept the natural process of matter evolution, accept the fact that the
Nature was the Creator, which had unlimited time, energy, materials, and other unknown,
but necessary conditions. The possibility of nucleation processes that characterize life on
the stage of geochemical and biochemical evolution of the Earth was proven in many stud-
ies (and a lot of authors refute it) A selections of publications on this subject can be found
in the monograph of G. P. Korotkova (1979) about the role of a “system of cycles”, con-
structive ideas contained in the works of V. I. Vernadsky (1930, 1964). The most con-
structive idea is that individuality was formed by the “biologization” of the cycle com-
ponents, which (components) were hitherto of a chemical nature. A new task — is aris-
ing: to find out what was the “attractivity of the biologization” for which the components
of the prebiotic cycle sacrificed their own (albeit relative) “immortality”?
The point is that the “immortality” is static by nature (because of restriction, and
possibly complete absence of adaptive abilities of the prebiotic cycle ingredients), it lim-
its the selection process and, therefore, the “job” of natural selection. “Biologization”
eliminated static nature of the cycles of matter and energy, enlivened the biosphere, cre-
ating conditions for the emergence of the biotic cycle of matter and energy, biological
evolution, escaping from entropy. There is another assumption: that biologization cycle
components could occur with the possibility of saving (extension) function of the com-
ponents which were biologized in the cycle. Therefore, the transfer mechanism of this
function already existed, and it could not raise up from anywhere but, from the preced-
ing (prebiotic) history of the components. I. e., along with the biologization of compo-
nents, the cycle and mechanisms of preservation and transfer functions in the cycle and
the biosphere were also biologized. Ontogenesis as a phenomenon was conceived in the
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bowels of the geochemical evolution and chemical cycles. Chemical reactions were
transformed in the morphogenesis.
Ontogenesis as a phenomenon encompasses all forms of life, ensuring existence of
hundred of times longer than the existence of their representatives: individuals. This is
one of the essential qualities of ontogenesis.
However, the presence of the existing diversity of life forms suggests there were sev-
eral ways of the evolution of ontogenesis, often calle several onthogenesis types in the
literature. Based on the fact that the main function (task) of the ontogenesis as a phe-
nomenon, in all types of life forms in general, is saving of the continuity of the species,
regardless the individuality life span. However, other properties and characteristics of the
ontogenesis as phenomenon may vary significantly: the duration of the prenatal period,
the duration of life of individuals, types of fertilization and reproduction, development
of embryonic membranes, direct/indirect types of ontogenesis, care for offspring, or its
absence, etc. These differences reflect the different evolutionary paths of ontogenesis, the
way gone to stabilize its basic and common properties of all the life forms (reproduction,
regardless of life span duration of an individual).
Further evolution of the ontogenesis seems to be done by the reorganization (evo-
lution) of mechanisms of morphogenesis, and restructuring of the ontogenesis process as
its general properties are stabilized. Compared to variety of specific ontogeneses (even if
you take the taxa of one class, for instance, mammals), the diversity of techniques or mech-
anisms of morphogenesis is minimal: division, differentiation, growth, migration (cells),
concentration, transformation, gastrulation, and perhaps a few others. But almost all of
these modes or mechanisms of morphogenesis are also highly variable. Variety of specif-
ic or particular ontogenesess is resulted from rearrangements (recombinations) and labil-
ity of morphogenesis modes that modify the course of ontogenesis in a particular envi-
ronment (and adapt the ontogenesis for changing environmental conditions.)
Based on the above, we have suggested that the evolution of ontogenesis as a phe-
nomenon ended with the emergence of life forms and types of high-level taxa (classes,
orders...), as well as general (common) features of ontogenesis. The variety of a partic-
ular ontogenesis is the result or consequence of adaptive radiation of species and mod-
ification of individual development by restructuring and lability of morphogeneses. This
issue we consider with the example of ontogeneses of the nestling and precocial birds
(Kovtun, Shatkovskaya, 2011): the new norm (“nestling”) is created based on the
“bird” ontogenesis is not a separate (nestling, half-nestling) type of evolution, but a mod-
ification of the basic “bird” type of ontogenesis. The more so that the stabilization of the
new norm is not the straightforward fact. There is evidence that a transition from the
nestlings to the precocial, and vice versa (Starck, Ricklefs, 1998), but, as is commonly
believed (the Dollo’s rule), the evolution is irreversible.
Summarizing this all, we consider the essence of ontogenesis (at the level of nowa-
days knowledge) to be providing the life forms with all levels of the organization, ability
and opportunity to produce “their own kind”, thereby maintaining the stability of the bio-
sphere and the dynamic cycle of matter and energy flows, i. e., procreation, and ability to
accept relay-race from predecessors to maintain their usual role and function in the
coenoses, biosphere and system cycles. This property is strictly controlled by the cycles of
matter and energy, through the natural selection, by selecting a normal phenotype, it also
selects the useful ontogeneses (Shishkin, 1981). Stability of ontogenesis is derived from here.
These essential features and objectives the ontogenesis is implementing through mor-
phogeneses, which accomplish the whole process of development.
The essence of the process (accomplishment of ontogenesis) is, apparently, in the
lability of its mechanisms, and the ability to direct the implementation of ontogenesis,
depending on the condition and needs of internal and external environment, provide adapt-
ability of ontogenesis stages, implementation of ontogenesis as a whole process.
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These definitions lack “internal relations” that characterize the essence of the phe-
nomenon and the process. Their interpretation, for nowadays is one of the main objec-
tives of biology of individual development.
The doctrine of ontogenesis (as creation of the theory of ontogenesis) seems to be
in some crisis: the lack of new ideas, vague (and sometimes absence) of so-called “orga-
nizing concept“. An idea occurs (perhaps quite an unpopular one) that methodological
constructivity should bring into the doctrine of ontogenesis certain, shall we say, “mod-
erate reductionism”.
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Received 3 January 2013
Accepted 21 March 2013
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