Dieser Aufsatz gibt einen Vortrag wieder, der 1994 an der Universität Bielefeld gehalten wurde. Er hat seine Aktualität nicht verloren.
The predominance of the organismic construction over physiological and genetical mecha-nisms
Karl Edlinger
Naturhistorisches Museum Wien, 3. Zool. Abt. Burgring 7, A-1014 Wien
1. Abstract
It can easily be shown, that science must be founded on a solid theoretical basis. This basis is achieved by a fundamenal analysis of acts (Handlungen) and above all by a succesion of acts, performed by a craftsman or a scientist. Each succession of acts enacts a theoretical assumption, a hypothesis about the nature of the object of handling.
From this presupposition physicians handle living organisms. If they keep up their living functions, they must conceive of organisms as hydraulic and mechanic systems, containing other subsystems which are partially of chemical nature.
In accordance with this approach the theory of organismic constructions configures organisms as hydraulic systems, consisting of a mechanical frameworks and a fluid filling. Form and organisation are enforced by a complex system of elements, acting in an antagonistic way. This framework comprises additional mechanical and chemical mechanisms. The action of all of these are highly coordinated and function in accordance to the requirements of the system as a functioning whole.
This concept is in accordance with the constructional demands of the mechanical framework, which allows only a certain order of procedures. As shown by construtvistic philosophers of science the results of empirical science are compatible with this view. The same methodological procedures should apply to biological research but are not yet worked out in a sufficient way.
2.Introduction
The biological disciplines show us a curious contradiction, which sets an important distinct mark between biology on one hand and astronomy, physics, chemistry, mineralogy, and geology on the other. The development of the non-biological sciences gave rise to a high level of mathematical sophisication and physical explanation. They were seen under the perspective of experimental testing and reliable prediction. In contrast the biological disciplines show this trend in a restricted way and only in such sections which are overlapping with other sciences, especially physics and chemistry. However, the very subject of biology, the living organism as a functioning whole, is not taken into consideration, it ludes the access of usual scientific procedures.
Attempts were made to solve this problem by formulating new models and theories on the basis of thermodynamics [1], synergetics [2, 3, 4, 5, 6, 7, 8], systems theory [3], the theory of chaos [9]or, last but not least, on a new variant of vitalism, called holism. However these models and theories are helpful for explaining processes of physics and technology and are of a general importance for a multitude of natural phenomena, but in no way specific for organisms. Systems theory refers to interactions and interdependencies of complex entities in general. It does not provide acceptable definitions of organisms and the special features of living beings. Holism is also not acceptable because it must postulate forces and mechanisms which cannot be studied by the methods of natural sciences.
3.Historical approach
Since I. KANT [10] reflected on „an eventually coming NEWTON of the blade of grass“, the problem of living beings has been permanently discussed. This philosopher hoped that a „NEWTON of biology“ might appear who would be able to point out the laws of the organismicity. These laws were expected to be inaccessible to the methods of Newtonian physics. Kant offered a temporary solution for this problem, in the form of an Aristotelian-teleological concept, which referred to the obvious autonomy, spontaneity, and intentional behavior of living beings but stood in cotrast to the simple causality of Newtonian physics. This dilemma continued to dominate biological theorizing till our days.
Even the philosophical endevour and the natural sciences of the romantic period could not solve the problem of organimiscity and develop an organismic concept of nature. During the romantic period the Newtonian sciences were substituted by a new holistic and organismic view of nature [11]. Biologists created new ideas of the characters of organisms, especially their autonomy, spontaneity and the capability of self-reproduction. But they failed to offer theoretical tenets which were in accordance with the well founded methods of classical Newtonian physics and chemistry. Newtonian physics continued to be accepted as the overruling paradigm.
4.Darwinism and the Synthetic Theory
A new situation arose when in the middle of the 19th century Darwinism was established as the predominant theory of evolution. Both, Darwinism and the Synthetic Theory as a subsequent stage of the theoretical development, ignore the organisms as specifically constituted entities. Darwinian theories are restricted to an extremely reductivistic view of life and its functions.
The exclusion of the organisms from reductivistic concepts is liable to ignore the difficulties and the problems which are generated by the observeable undisputable organismic properties. Only the particular aspects accessible to the reductionistic approach can be objects of investigation. The sum of particulate processes are easily confounded with the organismic whole. This characterization applies to the methodolgy and theorizing in genetics and molecular biology. The reductivistic methods which use physical and chemical procedures are helpful for the elucidation of the molecular and physiological mechanisms in organisms. Many of the molecular and physiological mechanisms worked out can be simulated and demonstrated in the laboratory.
As a consequence the Evolutionary Synthesis [12, 13, 14, 15] claims to be the authentic scientific interpretation of life, living organization, and evolution. Its proponents consider themselves to be the only users of correct and adequate scientific methods. Some of them pretend that reductionistic and darwinian views provide the basic tenets of a general philosophy of nature [16].
To bridge the gap between the simplicity of biochemical and molecular mechanisms and organismic properties the notion of information is used. Thereby it is pretended that complexity of organization might be reduceable to simplistic rules. The sequence of a few moluclar building elements is considered to be responsible for the complexity of organisation.
The chain of of amino acids in their dependence on the nucleotide sequences is conducive to the formulation of analogies between biochemical structures on the one hand and letters or words on the other. So it is not astonishing if reductionists believe that they have found the essential structures of organisms within the nucleus and the translating mechanisms of the genetic apparatus. So the genotype is pretended to be totally representative of the organism itself.
In this view, genes are thought to be interacting elements and organized as networks of hierarchic systems [17]. The structures beyond the genetic apparatus which is supposed to contain the information of the organismic whole are considered to be of minor importance or no real importance at all. The organisms lead a ghost-like life. However even in the perspective of reductionism the integration of the genetic mechanisms in the living entity cannot be ignored It is too obvious that genes can only funciton over the structure of the organisms. The outer framework for which specific principles and laws are not given are called phenotype. The phenotype is thought to consist of the sum of gene-dependent structures which are built by genetic informations.
Darwinism tries to convince a benevolet audience that the phenotypes are exposed to the selective influences of the environment [18, 19]. Under selective pressure some variants will survive and others will succumb. In this view the selective influence on the geneticic level is indirectly constituted over the dependence of the phenotype on the gneotype. In the last resort the genotype is selected from the environment. Again the organism and its specific properties are ignored, the reductionistic view prevails. Only the genotypus finds explicit recognition. The environment gains a determining influence on the molecular mechanisms and thereby on living organization. The concept is specific in describing an external selective influence on the genetic apparatus. The linkeage function of environment and genetic apparatus results from a very reduced view of the phenotype and of the organism as well. The term „organismic“ is only an empty play of words.
Thus, organisms seem to be well defined by the the genotypus. Biological fitness, as the most important idea and as the driving mechanism in Darwinian concepts of evolution presents evolution as process resulting from hereditary disposition, given by the genetic structures, in relation to external environmental mechanisms. This theory based dissolution and liquidation of the organism, commited by the proponents of „Synthesis“, continues, when evolution is defined in the terms of population dynamics. Evolution, in this view, consists of changes of gene-frequencies over time. The pressure of selection results in „well adapted“ genotypes characterized by gene-frequencies within the populations.
5.Models of origin and the early evolution of life
These ideas of molecular reductionism have a decisive influence on models for the origin of life [20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30]. Most of these models are focussed upon the emergence of the genetic apparatus and ignore the constitution of all other basic structures of organisms including those responsible for the functioning of the chemical and physical mechnisms described on the molecular level of the genetic apparatus. Organic substances like amino acids, nucleotides or lipids can be synthesized by technical arrangements in experimental machineries, which seem to simulate the conditions of the early time of earth. It is also possible to observe alterations of polymerized organic substances, which are self-reproducing with different speed and intensity. The rate of self-reproducing activity depends on the conditions constituted by the apparatus. The continous input of energy and material and the removal of the waste material are indispensible for the continuation of all of these processes. On this basis it seems possible to reconstruct the early processes of spontaneuos generation, alteration and adaptation of organic substances as primer elements of living beeings. In addition „cooperation-like“ interactions between organic molecules could be set going. These artficial procedures provide the experimental foundation of the theory of the so-called hyper-cycles presented by M. EIGEN and his group.The apparatus figures as the early environment
6.Critizism of reductionism
During previous years this reductionistic views and methods were frequently critizised. It can be shown, that most of the models of population dynamics are hypothetical and dont concern natural and real entities. In respect to early chemical evolution B. VOLLMERT [31] argued, that the physical and chemical conditions of the early earth, called „primeval soup“, give no chance for the formation of long chains of polymerized RNA and DNA-molecules. Hydrolysis would immediately destroy them. Similarly self-reproduction, cooperation and enzymatic reactions are dissipated and rendered functionally infeasible. The conditions which give freedom to this thermodynamic tendency must be counteracted by sophisticated technical arrangements or by compartimentalizing and enclosing structures of protobiotic entities. These can delimit the space of molecular mechanisms and exert sufficient cortroll on the enclosed processes. All of these presuppositions cannot be found in a non-living nature.
As a reslt of critcal studies M. EIGEN [19] was forced to concede that the hypercyles could only evolve in enclosures and necessitate compartments for an unditurbed function of the chemical mechanisms.
These conclusions can be generalized. All physiological and genetical functions appear be strongly dependent on surrounding and supporting structures which are given either in the apparatus or in the structure and organismic set up of living beings. There is no reasonable way to reconstruct prebiotic and the early stages of protbiotic evolution by concentrating on meolecular mechanisms alone. One he has to deliberately consider the role of the living apparatus and its structure. Here the principles of hydraulics must be invoked.
So each theoretical and practical approach to the question of organization and evolution must regard organisms as systems, consisting of interdependent different structural elements. Physiological and genetical mechanisms are conceiveable only as integrated but not as the dominating parts of organisms.
If EIGEN and other authors want to reach a proper evaluation of their prebiotic models they will have to stop regarding the molecular mechanisms alone and have to take account of the structural aspecs as constitutive for evolutionary alteration.
It is necessary to consider that evolution is a change of a complex structural whole containing numerous highly ordered elements. Genes and enzymes belong to these. As a consequence, for EIGEN the apparatus, not the „evolving“ substances it contains, must play the role of the organism.
The theoretical deficiencies of the reductionstic approach did not go unnoticed. Some other, non-biological „sciences“ concepts tried to figure as the theoretical basis for the definition of organisms and provide premises for the explanation of their specific properties such as self-organization, spontaneity, autonomy and for the unsuppressable tendency to evolve.
However, none of these approaches, systems theory, thermodynamics of open systems, the theory of dissipative structures, synergetics, or chaos-theories, were able to present a useful model or consistent concept. They are conducive to different inconguous experimental experimental arrangements and models, demonstrating the inability of an special discipline of physics or chemistry to construe an adequate concept of living organization. As pointed out above, the relation of the hypotheses to living beeings is undefined.
In this situation it would be a mistake to fall back on inappropriate and old-fashioned stages of biological research. Criticism of the kind formulated here is fully applicable to the understanding of organisation as propagated by classical biological disciplines of comparative anatomy, morphology and systematics. It is deplorable but necessary to state that in these fields no reasonable concept of the organism and its activity could be developed, because the subject of investigation was and continues to be the dead organisms, the cadavres, depleted of its living functions. However, there can be no doubt that a reconstitution of organismic biology can only spring up and prosper on the basis of morphological knowledge. Organismic and constructional concepts will rejuvenate mophology.
7.Towards new foundations of organismic theory
For a really consistent model of the organisms we have, in the first step, to clarify the theoretical approach to the problem of organism and organization. The second step will we have to elucidate the solid foundation of our methodological procedures. In accordance with some constructivistic positions, especially with the „School of Erlangen“ we must call for a theory of scientific handling and manipulation. We have to be aware that usefull behaviour and handling are dependent on the precise sequence of operations treating the objects of envestigation. But each operational step which comprises a manipulation of an object presupposes a consistent hypothesis about the nature and constitution of this object. It is possible to argue, that theoretical presuppositions, i. e. hypotheses, and usefull manipulations or experiments are serving as necessary supplements.
For physics and euclidic goemetry consistent basic theorems gained from some technical, especiall grinding operations, were given [32, 33, 34, 35]. In the case of biology and organismic theory we cannot figure out any kind of such operations neither within description and comparison of by traditional morphology nor in the explanations of physiology nor, as pretended by several authors, in the practice of breeding and cultivating. Morphology is focussed on cadavres and artefacts, the same is true for physiology which uses preparations and artificially isolated partial models of organisms. Selection as performed by man in breeding domesticated anmials or plants treats complete living organisms. Manipulation is confined to positively influencing the reproduction of appropriate individuals. Thus, the breeder takes the basic properties of organisms as given; he may even be unaware of the presupposed indispensible properties of life. By ignoring but presupposing the autonomy of life he produces an „unnatural“, artificial situation, in which some essential aspects of life under natural conditions are eliminated and compensated. From this follows that selection procedures and cultivating techniques are in no way sufficient to serve as premises for the constitution of living organisms. Darwinism is a basically misconceived access to living organisation and is in no way appropriate to justify the assumption of evolution.
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Medicine
Only medicine in its basic handling of patients provides the basic procedural requirements on which organismic concepts might be founded [36]. Rothschuh [37] as a physician developed a specific organismic theory which ha no counterpart in the field of biology. It is based on the experience of a medical understanding of the organism.
Interventions of physicans have to maintain and to support living functions of organisms. Medical descriptions of the living beings and their functions are mostly formulated in other terms than those considered appropriate in biology. Technical models and technic-analogous terms are of utmost importance. Most organismic functions can be described in the jargon of machine building and according to concepts of engineering. In many instances machines and machnie-like devices are used during life-saving medical interventions and treatment of diseases. This can be usefull only, if the machines function as real supplements of organisms and if the physican takes account of specific machinelike features of the organisms. Operational closedness of the body and of specific organs have to be regarded: Measurement Blood-pressure and other body fluids are essential indicators which can only properly evaluated in the frame of a constructinal understanding. The intactness of body cavity and the circulatory system have to be ensured. The turgescence and the hydraulic pressure of the tissues and cells, the character of the heart as a pump and the mechanical construction of the locomotion apparatus are objects of diagnostic evaluation. The possibility of utilization of energyconversion, the unimpaired and frictionless functioning of all deforming structures, the necessarity of harmonic activity of all mechanical elements, mechanical coherence, the cybernetic functions of the nervous system and so on.
Only after the dure acceptance of the aforementioned aspects can physiological and biochemical mechanisms taken into consideration. It is possible to monitor chemical and physiological processes of the living functions.
Only on the basis of preconceived constructional knowledge it can be legitimate to argue about gene-expression and other molecular processes. All of them depend on a preestablished structural and organisational constituion of the living entities.
Iin the case of experiments the structural and organsiational conditions are given by the set up of the apparatus, in the case of organisms, by the scaffolds, channels and pumping systems of the cells, tissues, organs and constructional wholes. It is obvious that the models of technology and engineering are fully applicable for organisms.
Chemical mechanisms are indispensable for the converting of material and energy in the metabolism. Enzymes are responsible for the metabolism, Their structure is dependent of the the DNA. In a similar way all proteins building up scaffolds of the cells and special tissues depend on the genetical machinery of the nucleus. But although the heredity of protein structures is accepted and well established in the biological sciences, we must consider the role of the DNA as an integrated part of the organism and not as the organism itself.
In this view organisms in general figure as hydraulic systems functioning like machines. It is hard to imagine what would happen, if physicians would try to save the life of a severely wounded person on the basis of self-organisation theories, systems-theories synergetics and thermodynamics or other models oulined above. Only adequate presumptions make us equal to the situation in this case, but this demand must not be seen as a postulate for a naive realism. It leads towards consistent theoretical organismic positions, which have to be proved by practice and corroborated by experiments. From these positions we can set up a model of the organism, which conforms to that of the theory of organismic constructions as presented by the Frankfurt group.
9.The theory of organismic constructions
This theory concieves organisms as hydraulic systems [38, 39, 40, 41, 42, 43, 44]. Hydraulic systems are under pressure of the fluid filling, which is surrounded by dense and flexible membranes. This causes a tendency to assume a globular shape. All deviations from this rule, i. e. non globular shapes of organisms or of their parts are enforced by tethering fibers, by surrounding packing rings, also consisting of fibers or by hard skeletal elements, secreted by glands or glandlike structues. Organisms are converters of energy and matter, which are deformed permanently by shortening of fibers, by energy-consuming gliding of actin-mysin-complexes. They can regain their lenght only in a passive way. Fibers are working in an anatagonistic way against other fibers or against the fluid filling and its surrounding membranes. Permanent deformations are generated by these mechanisms which cause the injection of energy into the environment. Contraction of fibers is elicited by stimulations, produced by pacemaker systems [44]. Contraction and restitutive expansio must be coordinated in a high degree. This is possible, because fibers and also microtubules composed of proteins form scaffolds, which have an additional effect in the enforcement of form.
The cellular and extracellular scaffolds can also be used for anchoring of contractile fibers. Enzymes can bind to them and compartimentation, as a primary condition of an effective metbolism, is achieved. So we can conclude, that organisms are highly ordered arrangements of mechanical and chemical elements, which are strictly adapted to their function within the organisms [45]. Mechanical structures function as frameworks for all other parts and components of the living machinery. This mode of internal „adaptation“ is the only one, which can be legitimately discerned in the kingdom of organisms. In the frame of the organismic machinery genetical and physiological mechanisms depend on the mechnical framework, i. e. the organismic construction. They are functioning as parts of its construction, supporting its activities and its permanent self-regeneration.
All reproduction activities must be seen as an aspect of energy conversion in the construction. The production of spermtazoans and eggs consists of energy driven formation processes. It results in the formation of separate mechanical constructions, which are able to develop into complex organisms. Ontogenetical development is also an energy driven process in mechanical constructions that is strictly guided by mechanical constraints. The differentiation of cells and tissues and the constitution of body architecture is only conceived as enforced by mechanical stress.
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New results and corroborations
Altough the reductionistic view of the Synthetic Theory is still dominating new results are helpful in confirming the biomechanic theory of organismic constructions. A growing number of authors demonstrate the predominant role of the compartimentation [46, 47, 48, 49, 50] and the indispensible and constitutive influences of biomechanics on biochemistry and molecular mechanisms [50]. Contractile fibers are capable of opening or closing ion-channels. Ions may exert a considerable influence on biochemical and molecular mechanisms. As shown by INGBER and other autors, deforming activity caused by cellular motility can alter the stiffness of the cytoskeleton. which may result in the alteration of the physical and chemical properties of the protoplasm. There is also evidence that shearing forces [51] are of great importance for the activation of some genes , and for the generation of mechanical stress that becomes in the differentiation of muscle cells and other tissues [52, 53, 54, 55, 56, 57, 58, 59, 60, 61, 62, 63, 64, 65, 66].
Mechanical force exerted by the extracellular matrix of metazoans is transmitted as signalling influence through the cell-mebranes and gains effectivity in gene-expression. As shown, epithelial tissue can be seen as induced by the self-arrangement of fibers and other material adjacent to a membrane and of the resulting mechanical forces. The extracellular matrix and the cytoskeleton can be influenced by stretching forces, and can be mutually effective and may restructure subsequent structures. Also mechanical forces could be shown to deform the chromosomal structures, when fibers functioning as actuating structures.
Lastly, at the level of organic molecules, the distinctness of biomchanics and biochemistry disappears [50]. This can be demonstrated by the presence of „mechanoencymes“, which are simultaneously functioning in molecular and mechanical mechanisms [47]. Ontogenetic development can be conceived as internal differentiation of mechanical constructions which enclose complicated chemical apparatuses. But the dominating structure in ontogeny, which becomes effective by constraining and enforcing the differentiation of cells, tissues, and organs, consists of the inner structural scaffold.
From this follows that cells must not be seen as separate and isolated units, building up the organism by cristal-like arrangement; in reality they are integral components of constructional frames.
Some of this results are the basis of the tensegrity model of INGBER and his group [52, 53, 55]. The tensegrity model gives reasons for the alteration of some features of the cell-construction, as is the stiffniss, by some deformation-activities. It is focussed on the laticelike inner scaffold of the cells, the cytoskeleton. The architecture of the cytoskeleton is held responsible for a manyfold of chemical processes.
This model may be seen as a progress towards a consistent theoretical basis for understanding living beings. At the other side, in its present form, it lacks the universality of application. The tensegrity model does not consider the role of hydraulics and the function of the different kinds of fibers for shape enforcement of cells, tissues and the organisms as wholes, which is given by the theory of organismic constructions. This theory shows us the interdependence of all levels and of all parts of living beings, functioning mechanically or chemically, and the fact, that organisms are ruled by mechanical principles, which influence all levels.
So we can conclude, that the distribution of various substances and genetical products, called „morphogens“ [67, 68, 69, 70] may result out of differentiation, but they are not the primary cause of this processes.
11.Autonomy of organismic constructions
The mechanical energy driven framework of the organism has, in the case of animals, a great deformability, which is becomes effective in motility, propulsion, capture of food, expulsion of waste, and blood circulation. Locomotory deformations can display a variety of modes in dependence on the working construction. Motoric patterns are usefull, if there is no conflict between the actions of different components. Similar to some machines, the propulsive apparatus can be used in different ways. The only demand is, that deformations remain adequate to the constraints given by the organismic construction. The different deformations must be harmonized in a high degree. A manifold options of locomotion become manifest in a high degree of organismic autonomy. Autonomy as resulting from energy driven activity of constructions is not consistent with the basic tenets of molecular reductionism and totally at variance with the Sythetic Theory of evolution.
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Relations with the environment
If we accept the hydraulic and mechanic nature of living beeings, the interdependence of all organismic levels and the predominance of the organismic construction is apparent. The living being has the form of a well defined apparatus which is moved by its own intrinsically generated deformations on the basis of energy consumption and under the influence of internal pacemakers. Actions of organisms have a predominant internal aspect. All actions are performed in relation to external and environmental factors [71, 72]. The constitution of the organismic construction and its mode of functioning determines the external „contacts“ and the organismic interdependencies with the environment. The environment can only be actively conquered by the organisms. Environmental factors must never be understood as externally generated forces or constraints working on the organismic constructions. There is no adaptation generating influence of the environment [73]. The constructional properties of the organismic units and their mode of working are responsible for survival, reproduction and even the death of organisms in their habitats. There can be no greater contradistinction to the the synthetic theory.
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Conclusions
A consistent definition of the living organism as the object of biology cannot be derived from reductionistic concepts of molecular biology and the Synthetic Theory of evolution which nowadays dominate the whole field of biology. The presuppositions of these all-encomapassing theories are not sound. Explanatory theories, in the view presented here, contain as essntial elements premises of handling of the respective objects. We must take into consideration, that each mode of handling must be done on the basis of a preestablished theory, which clearly refers to the most important features of the object. This approach can muster knowledge and procedural strategies of medicine, a science, which can prove its success and explain its failure in saving life and treating living functions.
On these premises organisms must be configured on a new paradigm, conflicting with the Synthetic Theory and the extremely restrictive expectations of reductionisms. In organisms the mechanical constructional frame and its internal coherent structure predominate all other levels and particulate molecular and physiological mechanisms. In this view organisms are highly ordered mechanical systems which are organized according to mechanical and chemical laws subservient to the requirements of survivial. Within living beings genetics figures as very important but not as the only life determining mechanisms.
During ontogenetical development differentiations of cells and tissues follow of mechanical constraints, generated by the construction of the organism. Organisms are autonomous and establish external environmental relationships according to their internal constitution. Consequently environmental factor have to be construed as only determined by the Properties of the organismic constructions. So the idea of adaptation, the suggestion that organisation might be ruled by environmental factors is reversed and eliminated from organismic biology. So organisms in general appear as autonomous and not any copies of ecological conditions.
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