The Theory of Organism

Theory Biosci. (2000) 119: 276317 Urban & Fischer Verlag http://www.urbanfischer.
de/journals/theorybiosc
The Theory of Organism and the Culturalist Foundation of Biology

Mathias Gutmann1 & Eva M. Neumann-Held2
1 2
Institut fur Philosophie, Philipps-Universitat Marburg Europaische Akademie zur Erforschung von Folgen wissenschaftlich-technischer Entwicklungen GmbH, Wilhelmstr. 56, 53474 Bad Neuenahr-Ahrweiler
Address for correspondence: Mathias Gutmann, Institut fur Philosophie, Philipps Universitat Marburg, Blitzweg 16, 35039 Marburg Received: July 17, 2000; accepted: August 28, 2000 Key words: Organism, Function, Structure, Constructional Morphology, Culturalism
Summary: After the disappearance of organism was diagnosed, the discussion about the role of a theory of organism in biology is characterised by a significant contradiction. On the one hand, the importance of a theory of organism is stated. Particularly developmental biology demands organism-centred approaches as a basis for conceptual integration. On the other hand, several modern biological disciplines such as genetics and molecular biology simply don't need a theory of organism for their work. Consequently, the determination of the status of the organism and its relevance for biology at all is an unsolved problem. In order to clarify the methodological status of the organism in biology we start with the reconstruction of three important propositions. A life oriented approach and a hierarchy concept which both are from a neo-Darwinian origin are confronted with a structuralist approach of organism, that can be characterised as a non-Darwinist approach. Our own attempt for the solution of the organism problem applies the tools of culturalist methodology. In accordance to this pragmatic approach, the term organism is introduced as a concept of notion. A constructional morphological case study exemplifies the applicability of this concept. From the culturalist point of view a methodological foundation of biology can be achieved, that provides a consistent basis for a comprehensive integration of biological knowledge.
1 Introduction
A clarification of the status and the nature of the organism has a significance that reaches far beyond the realm of biology alone. The organism has been and still is used as model and metaphor in the effort to describe relationships between parts and wholes in very different areas, to conceptualise organisation, differentiation, and even the social embeddedness of structures or parts of the entity they comprise.
The theory of organism and the culturalist foundation of biology
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Such so-called organicist approaches have a long history in Western philosophy, as do some typical features of organicist approaches, such as wholeness and structural closure (e. g. the naturalist justification of Platon's ideal state see Schafer 1993)1. In modern times Kant2 (1983) noted that the or ganisation of modern states could be compared to the relationship between parts of an organism. Such an analogy is obviously based on the idea that as in an organism, the parts in the organisation of a state depend functionally on each other, that they are mutually means and ends for each other at the same time3. Likewise, we find in romantic political concepts the term organism as a metaphor for human communities (see Taylor 1989 for further reading). Nowadays, organicist descriptions are frequently used in a variety of philosophical approaches. Examples are the theory and philosophy of culture (e. g. Sponger 1983), theory of action (see Mead's 1992 a & 1992 b, 1998 and Peirce's 1996 application of biological knowledge) and even linguistic or hermeneutic approaches (see Humboldt's 1985 usage of the word organism and organisation for language; see Gadamer 1990). The same is true for the approaches of philosophical anthropology that usually refer (directly or indirectly) to a theory of organism (see e. g. Gehlen 1986, Plessner 1975). Even within the context of theory of arts, the term organism seems to be applicable (see Eggebrecht 1986). In addition to these examples from philosophy, theory of culture and arts, organicist positions are also prominent in politics, sociology and history. The organism obviously plays an important role in the construction of theories of numerous disciplines. This presumes that the status and the nature of the organism are settled. Here, it is usually assumed that the clarification and definition of the organism is the typical project and even the object of biology, and that therefore organicist positions are securely grounded in biological knowledge. It must come as an unpleasant surprise to realise that in modern biology, the
For a discussion of such features see section 4. As the term organicist is a diagnosis, based upon historical as well as systematic reconstruction, we will refer only to some features that are tightly connected with such approaches. As the term organism from ancient Greek time to nowadays changed its meaning fundamentally, the identification of an author as an organicist provides severe problems of comparability that can be avoided by the characterisation we applied here. 3 Kant writes: Man kann umgekehrt (because a direct analogy of the organisation of nature with any causality we know, the authors) einer gewissen Verbindung, die aber auch mehr in der Idee als in der Wirklichkeit angetroffen wird, durch eine Analogie mit den genannten unmittelbaren Naturzwecken Licht geben. So hat man sich, bei einer neuerlich unternommen ganzlichen Umbildung eines groen Volkes zu einem Staat, des Worts Organisation haufig fur die Einrichtung der Magistraturen u. s. w. und selbst des ganzen Staatskorpers sehr schicklich bedient. Denn jedes Glied soll freilich in einem solchen Ganzen nicht blo Mittel, sondern zugleich auch Zweck, und, indem es zu der Moglichkeit des Ganzen mitwirkt, durch die Idee des Ganzen wiederum, seiner Stelle und Funktion nach, bestimmt sein. (Kant 1983: 323) Note here that Kant uses the German word Staatskorper, which means literally body of state.
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loss of the organism is almost a truism4. With the loss of the organism biology appears to have lost its foundation as well, and one might also wonder what this can mean considering that we are obviously surrounded by organisms, that is by plants, animals, and other living beings. Indeed, the word organism can be used exactly in the same sense as and as a synonym for living beings. If organism is used in the sense of living being then it seems that in some areas of biology the organism (the living being) no longer plays a significant role. In molecular biology, for example, some biochemical molecules might stem from living beings (although these molecules can be synthesised as well), but for most experimental approaches the living tissue is at best one of numerous other purely technical settings in which biochemical reactions occur. Organismically derived is not the same as reactions typically observed in living beings. A second example is evolutionary biology. Here, many approaches model evolution by focussing on the differential distribution of heritable patterns, thus viewing the organism itself as nothing but some kind of vehicle for the heritable entities that are assumed to be essential (e. g. Dawkins 1986 & 1992). On the other hand, in some areas of biology the organism continues to play a significant role and can be found in morphological approaches as well as within the context of ecological, developmental and evolutionary concepts (e. g. Driesch 1935, Duncker 1998, Goodwin 1990, Gutmann W. F. 1995, Roux 1881, Tansely 1935, Uexkull 1973, Wagner & Laubichler 2000, Webster and Goodwin 1982 & 1996; for further reading s. Weingarten 1993, Potthast 1999). Often, these approaches have a non or an anti-Darwinist impulse5, particularly when they use the word organism as a technical term, so that different approaches developed different meanings for organism. This situation becomes even more complicated when we consider that several of these approaches are of an explicitly non-biological, mostly philosophical origin, but nevertheless refer to or incorporate biological knowledge and theories to corroborate the intended speech about the organism. So, today we face a curious situation in which, on the one hand, the organism has lost its central place in modern biology, but on the other hand numerous different organism-centred approaches have been developed from a melange of biology and philosophy. This situation has prompted some philosophers to pose the question of whether the organism is a bio4 5
For the discovery of the disappearance of the organism see Webster & Goodwin (1982 & 1996), Goodwin (1997). We distinguish between the terms Darwinian and Darwinist. Darwinian refers to one type of evolutionary theory that can be gained by reconstructing Darwin's approach of breeding as a model for evolution. Darwinist approaches, for example mathematical population genetics, have systematically reduced the Darwinian approach. Interestingly, it can be shown that anti-Darwinist approaches may be in some respects even compatible with Darwin's own conceptualisation of the structural problems of a comprehensive evolutionary theory (for the detailed argument, see Gutmann 1996, Gutmann & Weingarten 1999).
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logical, a philosophical or even a philosophical-scientific problem (see, for example, Mahner & Bunge 1997). Rather than trying to answer this question, which in some ways presupposes the existence of (a) correct organism concept(s), we suggest taking a closer look at alternative approaches in biology. We will therefore start our analysis with a reconstruction of some applications of the word organism in biological theory. This will lead us to the conceptual origin of the loss of the organism. We will then contrast these results with an example of an alternative approach, which attempts to reestablish a theory of organism that gives the organism a central place in biology. Attempts of this kind come mainly from the field of developmental research6. Therefore, the next part of this paper presents an analysis of one of the most substantial approaches of this type, the structuralist approach. Using theoretical, philosophical, historical and biological perspectives, the structuralist approach attacks dichotomies, such as the phenotype genotype distinction, which have replaced the organism in the Darwinist approach (see, for example, Webster and Goodwin 1992 & 1996). Although anti-Darwinist and Darwinist approaches are antagonistic in several important aspects, we nevertheless think that both have severe shortcomings. In the last part of this article we will therefore present our own approach to the problem: a proposal for a theory of organism, based on an explicit consideration of the explanatory goals, the anticipated knowledge, and the status of the resulting explanations and descriptions. This proposal not only avoids the typical misunderstandings of current organism concepts but also provides a basis for the determination of the limits to the realm of biology itself.
2 The Organism in Modern Biology Its Loss and its Defence

Our investigation will begin with a re-visit to the loss of organism from biology. This requires a reconstruction of the main applications of the word organism, for which we will use two examples. Our first example delineates the loss of the organism through the identification of the organism with life. The second example will try to localise the organism in so-called natural hierarchies. This will lead to some more general reflections on the status of natural hierarchies and ontological assumptions in the construction of biological (scientific) theories.
6 Developmental biology was traditionally always more anti-Darwinian than other fields of biology (for a historical outline, see Jahn 1990, for a systematic argument Weingarten 1993).
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2.1
Organism Lost in Life
It might appear as a surprising starting point to begin our search for the loss of the organism with Mayr, for whom organicism is a characteristic feature of a specific conceptual tradition in modern biology. However, we will show that it is precisely in organicism of the type that Mayr characterised and which has been highly influential in the major parts of biology that the organism is lost as an entity with definite properties. In his efforts to establish biology as a science Mayr presents an instructive and revealing characterisations of organicism by emphasising two main points: the holistic and the emergentist aspect. Mayr writes:
To sum up, organicism is best characterised by the dual belief in the importance of considering the organism as a whole, and at the same time the firm conviction that this wholeness is not to be considered something mysteriously closed to analysis but that it should be studied and analysed by choosing the right level of analysis. (Mayr 1997: 20)
Therefore, Mayr seems to hold the view that the organism is just one particular level in a hierarchy of life. This level can be analysed, so Mayr continues, by referring to the relevant lower levels of this hierarchy:
At the molecular level, all and at the cellular level, most of their functions obey the laws of physics and chemistry. There is no residue that would require autonomous vitalist principles. Yet, organisms are fundamentally different from inert matter. They are hierarchically ordered systems with many emergent properties never found in inanimate matter; and, most importantly, their activities are governed by genetic programs containing historically acquired information, again something absent in inanimate nature. (Mayr 1997: 20 pp)
Note here Mayr's emphasis that organisms have to be clearly distinguished from inanimate matter after all, this seems a requirement for the determination of the object of biology itself. But organisms also have to be thought of as hierarchically ordered systems, in which each level can be analysed by chemical and physical means, and which obeys natural laws. Based on the fact that numerous levels of biological organisation can be distinguished we first started with the question: What exactly defines one of the levels in the hierarchically ordered system usually given as the level of organism? A second question, apparently connected to the first, then arises: How is life to be distinguished from inanimate matter? And clearly, Mayr seems to think that both questions can be resolved in one approach, which points out the characteristics of life. In his view they are obviously also the characteristics of the level of the organism:
These properties of living organisms give them a number of capacities not present in inanimate systems: A capacity for evolution, a capacity for self-replication, a capacity for growth and differentiation via a genetic program, a capacity for metabolism (the binding and releasing of energy) a capacity for self-regulation, to keep the complex system in steady state (homeostasis, feed-back), a capacity (through perception and sense organs) for response to stimuli from the environment, a capacity for change at two levels, that of the phenotype and that of the genotype. (Mayr 1997: 22)
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According to Mayr these features characterise a living organism. Organisms as multi-leveled hierarchically ordered systems can be distinguished from other kinds of such systems, because organisms are fundamentally different from inanimate matter. Therefore the next question is: what distinguishes inanimate matter from animate matter? Surprisingly, Mayr explicitly uses here the same properties as being those of life. One might argue now that the systematic problem with these defining properties is that combinations of them, or sometimes even all of them, could equally well be attributed to inanimate matter. Mayr's definition only seems adequate because he applies an implicit background knowledge that refers to a sufficient difference between animate and inanimate entities. Without this background knowledge the properties are not appropriate for distinguishing life from non-life. What does this mean for the definition of the organism? Note, that when Mayr gave the properties that distinguish animate from inanimate matter, he related these features not only to animate matter that is life but to living organisms as well. Since organisms were defined so far as multi-leveled systems which can be distinguished from other systems by belonging to animate matter, organisms have been introduced so far only as being nothing more than a part of animate matter. Therefore, when Mayr uses the expression living organism (see quote above) the extension organism is unnecessary or redundant. The term organism is reduced to a word that does not add any relevant aspect to the object of biology, which is life (animate matter) itself. What then are the specifics and the significance of organicism for biology? A further quote from Mayr is helpful here:
Since the 1920s, the term holism and organicism have been used interchangeably. Perhaps, at first, holism was more frequently used, and the adjective holistic is still useful today. But holism is not a strictly biological term, since many inanimate systems are also holistic, as Niels Bohr pointed out correctly. Therefore, in biology the more restricted term organicism is used now more frequently. It encompasses the recognition that the existence of a genetic program is an important feature of the new paradigm. (Mayr 1997: 17)
Indeed, the assumption that holistic systems exist in the realm of inanimate matter as well is true for many if not all the criteria, which Mayr applies in order to distinguish, animate from inanimate matter. Another most important point here is the significance given to the genetic program. The genetic program comes to be the most important criterion for the difference between inanimate and animate entities, as a further quote stresses even more:
Every system, every integron, loses some of its characteristics when taken apart, and many of the important interactions of components of an organism do not occur at the physicochemical level but at a higher level of integration. And finally, it is the genetic program, which controls the development and activities of the organic integrons that emerge at each successively higher level of integration. (Mayr 1997: 20)
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Obviously, the level of organism within the hierarchy of life (animate matter) is still not determined by applying any specified set of criteria. Instead, Mayr continues to analyse the difference between inanimate and animate matter, which he defines by the existence of a genetic program, or more precisely, by a particular dualism, the phenotype-genotype relationship. Mayr writes:
The dualism of modern biology is consistently physicochemical, and it arises from the fact that organisms possess both a genotype and a phenotype. The genotype, consisting of nucleic acids, requires for its understanding evolutionary explanations. The phenotype, constructed on the basis of the information provided by the genotype, and consisting of proteins, lipids, and other macromolecules, requires functional (proximate) explanations for its understanding. Such a duality is unknown in the inanimate world. (Mayr 1997: 21).
For Mayr, therefore, the genotype-phenotype dualism reflecting the genetic-program metaphor serves as the defining aspect of life. The organism, on the other hand, has been introduced only as multi-leveled system of animate matter. Since apparently the main characterisation of animate matter is the genotype/phenotype distinction, it has to be concluded that this is also the main feature of the organism. In other words: the organism as a specific structure in its own right does not hold a place within the hierarchy of life that could not equally well be expressed by referring to other descriptions of living entities (such as the genotype-phenotype). In Mayr's view the categorical difference between approaches which refer to the organism rather than to life as the object of biology, is denied by refusing to define life and organism differently. Reducing the speech about organisms to speech that deals with life as the very object of biology, the term organism remains a word only, and the dualism between life and organism collapses to a dualism between phenotype and genotype. At this point we can summarise the result of Mayr's approach: the organism, which was assumed to be a putative biological unit of unique character, is lost in its identification with life, which is distinguished from inanimate matter by the genetic program. The unity of biology is thereby resolved by creating an unbridgeable gap between phenotype and genotype7.
2.2
Organism Lost in Hierarchies The Schism of Biology
We started the above analysis of Mayr's approach with the aim of finding a definition of the particular level of organism. Since Mayr had demanded that the organism should be analysed by choosing the right level of analysis, we had posed the question which criteria would determine the level of
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The inclusion of emergence does not solve the methodological problem here, because emergence can be reconstructed as a confusion of different language levels (s. Gutmann 1999).
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the organism among other levels of analysis of animate matter (Mayr names, for example, the molecular and the cellular level, see quote above). However, although Mayr uses the word organism and even defends a particular kind of organicism, he nevertheless loses the organism in the gap between phenotype and genotype. Our search will now lead us to a second author, Niles Eldredge, in whose writings the organism plays a central role, this time not in one hierarchy but in two. The question will again be whether we can find any definition(s) that specifies (specify) the particular level of organism(s) in any of these hierarchies. First of all Eldredge claims a schism of biology, which is reflected in the fundamental difference between the realms of ecology and evolution. As a result of the gap between ecology and evolution, two hierarchies must be developed, which are separate fundamentally, intrinsically, and ontologically (Eldredge 1992: 3). Following this dualistic scheme, the units of the different hierarchies differ. Starting from the top, the genealogical hierarchy, made up of populations (= reproductive unites), consists of 1. monophyletic taxa, 2. species, 3. demes, 4. organisms. 5. chromosomes and 6. genes. (Eldredge 1985: 188) The ecological hierarchy, made up of the atavars (= economic units) consists of 1. regional biotas, 2. communities, 3. populations, 4. organisms, 5. cells and 6. molecules. (Eldredge 1985: 188) A first comparison reveals the term organism in both hierarchies at position 4. And although the two hierarchies are irreducible and therefore the roles played by organisms change, depending on the membership of hierarchy8, Eldredge states:
Organisms are simultaneously parts of both systems. In other words, any single organism (with some special exceptions) is always a part of a local economic system, as well as a local reproductive population. Because this is so, it is obviously that the two hierarchies converge at the organism level (. . .).(Eldredge 1992: 3).
This convergence of hierarchies plays a significant role in Eldredge's concept, because from here it is possible to consider putative relationships between both hierarchies, which again allows the processes in both to be conceptualised as participating together in the evolutionary process. In Eldredge's words:
8 Note here, that Eldredge's dualism is not in contrast to the phenotype-genotype gap, but the difference between germ lines and soma becomes significant below the organismic level, ensuring the irreducability of the two hierarchies not only above, but also below the level of organism. Eldredge writes: Below the organism level, of course, the distinction between the somatic and the germ lines (i. e., in multicellular organisms) once again ensures a clean separation of the elements of the two hierarchies. Hence the conclusion (Eldredge and Salthe 1984) that there must in fact be two independent, yet parallel and interacting, process hierarchies that together combine to yield evolution. (Eldredge 1985: 175)
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Hierarchy allows us to acknowledge that both ecological and genealogical entities, events, and processes are involved in the evolutionary process. The entities all appear to be stable individuals. They are hierarchically arranged. There are processes intrinsic to each level that are not reducible to those of lower levels subsumed by higher levels. (Eldredge 1985: 214)
For Eldredge, different kinds of relationships between these two hierarchies are imaginable, and he discusses two possible extreme positions that define a whole spectrum of combinations (Eldredge 1985: 187 pp): 1. There may be one single level, thought to be predominant (this may be the organism, but can of course also be any of the other levels). 2. The relationship between the reproductive and the economic hierarchy can be determined in terms of completely symmetrical mutual relationships between each single level of both hierarchies. Of course, the solution of the problem of how the two sides may be brought together again depends methodologically on the answer to the question of whether or not the elements of the two hierarchies are really comparable, i. e. whether or not the references of the respective entities are identical. And here we come back to the main problem that we are concerned with in this paper. The fact that in both hierarchies at the same place (given here as position 4) the same word organism can be found, does not imply that the same reference is meant. The problem we are facing is not only that it is not clear by which characteristics the level of organism is determined, but furthermore that there are two hierarchies in which the term organism might even have different meanings. This last point could only be excluded on the basis of strong ontological assumptions about the nature of nature. For the organism this has two alternative consequences. In the first case, the organism simply denotes things like plants and animals. Here the term is not part of a scientific language, and the process of its transformation into such a scientific (definable) term remains as unclear as its status in any of the presented hierarchies. In the second case the organism is specified, and its characteristics are given in scientific language. Following the hierarchy argument the characteristics are then given either by reference to a lower level (e. g. in terms of the genetic code or the constituting molecules) or to a higher level (e. g. in terms of economic or reproductive relations, demes, atavars etc.). In these cases the organism can be replaced by the respective descriptions. In all the cases that stem from the model of hierarchical description of nature or life reconstructed here, the organism is lost.
2.3
The Status of Hierarchies for the Foundation of Biology
So far, it has been shown that the organism is lost in the framework of the hierarchies of nature or of life (animate matter) that have been discussed. We might now ask, how are the proposed hierarchies themselves justified? Therefore, in this section we will work on a more general aspect, namely the argumentative function and the methodological status of hier-
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archies in biology. This investigation is important in view of our aim to introduce our own approach towards a concept of an organism. For this purpose it is useful to be clear whether this must be preceded by the construction or the acknowledgement of a hierarchy of nature. After all, the two approaches considered above showed that hierarchies usually serve as a background ontology, which shall provide what could be called the realm of biology. The organism would then have to be found if at all as part of such a naturally given hierarchy. In ontological approaches the determination of the status of the organism and the determination of the realm of biology itself is thus closely linked to the natural givenness of the hierarchy of nature. Therefore, it is necessary to investigate this claim of the ontological foundation of biology. Mahner & Bunge (1997) have very explicitly and in sophisticated philosophical terms explored the prerequisites of an ontological foundation, which is at the basis of many approaches to biology (and the sciences in general). Therefore, we will concentrate on their work to describe the ontological prerequisites and to state its limits. In an ontological foundation the realm of biology is usually given in terms of description, i. e. scientific theories are thought to describe the world at least in parts. From this point of view an ontological foundation (in general) is not an optional but a necessary prerequisite for every single science, stated in the words of Mahner & Bunge as follows:
If ontology is general science, then the specific factual sciences, or sciences of reality, are special metaphysical or regional ontologies. In our view, both sciences and ontology inquire into the nature of things, but whereas science does it in detail and thus produces theories open to empirical scrutiny, ontology is extremely general and can be checked solely by its coherence with science. (Mahner & Bunge 1997: 4)
The relation between science and ontology is described here in a Husserlian manner as a first science, from which constitutive elements of the factual or single sciences of reality are derived as detailed research approaches. From this point of view ontology is indeed a first science, which gives the very general structures of the nature of things, which themselves are subjects of inquiry by ontology as well as by sciences. Consequently, some very general propositions (called axioms or postulates) can be given, which describe the references of scientific theories in a sequence of decreasing generality. According to Bunge & Mahner:
Postulate 1.1. The world (or universe) exists on its own (i. e. whether or not there are inquirers). (Mahner & Bunge 1997: 5)
As to the status of postulate 1.1, they state:

This axiom is not provable, and it tells us nothing about whether the world can be known and, if so, to what extent. (Mahner & Bunge 1997: 5)
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And furthermore:
A philosophical principle shared by most philosophical schools is that all things are in flux. (To be sure, although this principle has never been refuted, it is hard to prove: this is why we must postulate it.) Thus we introduce: Postulate 1.5: Everything changes. (Mahner & Bunge 1997: 17 pp).
If we introduced the realm of biology in terms of postulates, that give the ontological characteristics of the respective objects, then a classical hierarchy of levels, called biolevels, can be defined. The investigations of the former chapters, however, have led to the question: How can such (any) hierarchy be justified? Clearly, the justification of such hierarchies requires a closer look at the methodological status of the postulates, from which the respective definitions (e. g. the definitions of the single biolevels) are derived. The postulates named here are obviously very general; and it is stated (correctly) that they are not falsifiable. If ontology is indeed a necessary prerequisite of sciences9, the quest for an alternative must stop here. However, if it could be shown that accepting ontology as a prerequisite of science leads to methodological problems, it might be advisable to examine the possibility that ontological assumptions might be neither necessary nor useful for the construction of scientific theories. Indeed, when we take a closer look at the relationship between ontology and science, we can detect a significant dilemma. On the one hand, ontology is seen as a field of fundamentals that are needed for sciences (described as the field of regional ontologies). But on the other hand, it is almost always possible to suggest different propositions for ontologies. Therefore, we need criteria in order to determine the correct ontology10. And here is the dilemma, which easily becomes a vicious circle: If coherence with sciences is thought to be relevant for the selection of the correct ontological basis, this basis cannot be constitutive for the sciences themselves. If, alternatively, the sciences of reality give the criteria for the correct ontology without forming a vicious circle, then the sciences must be possible without an ontological foundation of the proposed type. In that case, however, we are in the difficult situation that independently of an ontological foundation criteria for the correctness of the sciences and their assumptions must be found. The fundamental problems we are facing in defence of ontological statements of the proposed kind can be exemplified in the cases of the hierarchies of nature that have been discussed. Distinct propo9
This is a weak implication of the assumption that the relationship between ontology and sciences is the relation between ontology and regional ontologies, whereby the latter is the field of the sciences of reality. 10 Here, it is often suggested that the criteria for the right ontology are its coherence with science. As a model of such coherence approaches one could think of the Duhem-QuineThesis; on Holism from an epistemological point of view and for further reading, s. Stegmul ler (1987).
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sitions of hierarchies can be made, which are simply irreducible (See Tab. 1 and 2).
Table 1. The table gives an overview of several hierarchies that can be found. Note the differing structure of these hierarchies as well as the different levels that are mentioned. Ecological (Odum 1991: 39)11 Biosphere Biogeographical Region Biom Landscape Ecosystem Living community Population Organism Physiological (Odum 1991: 39) Individual Organ system Organ Tissue Cell Organelle Molecule Genealogical hierarchy (Eldredge 1985: 187) Monophyletic taxa Species Demes Organisms Chromosomes Genes Ecological hierarchy (Eldredge 1985: 187) Regional biotas Communities Populations Organisms Cells Molecules
Table 2. The table continues the overview of Table 1 Bio-levels (Mahner & Bunge 1997: 178) B 1 = elementary cells = set of all elementary cells B 2 = composite cell level = the set of all composite cells B 3 = organ level = the set of all (multicellular) organs B 4 = multicellular organismic level = the set of all multicellular organisms B 5 = population level = the set of all biopopulations B 6 = community level = the set of all communities (biocoenoses) B 7 = biosphere level = the set of all biospheres Vertical aspects (Solbrig 1994: 4312) Molecules Cells Organs Horizontal aspects (Solbrig 1994: 43) Populations, species, living communities
The hierarchies in Table 1 and 2 clearly differ in several aspects, especially in the number of levels, their names and their relationships. The last hierarchy is a particularly good example of a mixed concept, because there are only three levels in the vertical direction and three further units of nature, which are thought to refer to each of these levels and to each other. With these examples in mind the methodological problems identified by discussing the standard ontology of Mahner & Bunge (1997) can be demonstrated. If distinct hierarchies are possible (at least in the sense of thinkable), then there should be some criterion that allows the correct one to be identified. Alternatively a principle or rule of translation or correspondence must be stated, which allows one or all the single levels of these hierarchies to be correlated.
11 12
Translated from the German edition. Translated from the German edition.
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But even if this were possible, there still remains one fundamental difficulty to be solved by all concepts of hierarchies of nature: the explication of any principle for the construction of a hierarchy. Only such a principle or rule evaluates whether or not a proposed hierarchy was properly constructed, at least in terms of its completeness and adequacy. At this point of our attempt to identify the place of the organism in the realm of biology, hierarchies have not been particularly helpful. Although we cannot show it in detail in this paper the same holds true if the realm of biology is determined ontologically not by reference to hierarchies but by alternative metaphors, such as a multitude of processes or network.
3 Introducing the Organism as a Unit of Transformation

The attempts to determine the relevant features of an organism in terms of a list of putative characters of life or a natural hierarchy has lead to some severe methodological objections, as was shown in the preceding sections. Setting aside the differences in detail, the approaches discussed (Mayr and Eldredge) show at least one fundamental similarity: They refer to a specific concept of evolution, which includes the fundamental gaps we have noted between genotype and phenotype and between ecology and evolution. On such an evolutionary basis the organism and its proposed role in biology is considered very strictly in terms of the explanatory goals of evolutionary theory. This in itself is not problematic, but becomes so when it is assumed that an evolutionary description of the organism is sufficient for all purposes. This would imply that the theory of evolution is indeed the comprehensive and fundamental biological master program, as it was characterised for example by Dobzhansky in his famous quote: Nothing in biology makes sense except in the light of evolution (Dobzhansky et al. 1977). However, following the design of such a master program results, as we have seen, in the loss of the organism as a relevant scientific entity. This is exactly the result which Webster and Goodwin derive as well:
Organisms have disappeared as real entities from biology. Their essences are identified with hereditary determinants, which Weismann located in a distinct part of the egg, the germ plasm. The information in these determinants, now identified with DNA sequences in the chromosomes, directs the development of the visible body of the organism from the rest of the egg substance, the somatoplasm. (. . .) Weismann's dualism saved Darwinism from inconsistency by excluding the possibility of Lamarckian inheritance, and it provided a clear rationale for studying inheritance independently of development, since embryogenesis can be reduced to the activities of genes as the primary determinants of morphogenesis. (Webster & Goodwin 1996: 131)
In contrast to this Darwinist position and its constitutive dualism, the structuralist approach is formulated as an alternative proposing a theory of organism in which the organism remains a real and constitutive entity of
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biology. It is a morphogenetic approach and centres on the form. To understand what follows it is necessary to realise that although Webster & Goodwin ask for the return of the organism in biology, the term organism stands at first glance more or less for the living being and is therefore a non-scientific term that simply denotes plants and animals. Introducing the relevant scientific (technical) term form, however, integrates the organism into scientific theory (for a reconstruction of this program s. Gutmann & Voss 1995). The form must not be misunderstood as an empirically evident matter of fact description, as it might be in the context of classical or traditional morphology. Therefore, the form is not the organism (that is, the living being) or the empirical outer shape of the organism (the living being) or its parts. Rather: the form is a mathematical representation of generative relations that refer to the whole or, as will be relevant later, to parts of organisms:
In the theoretical models proposed by Goodwin, the field is conceived as a dynamical system and he argues that genetic and environmental factors determine parametric values in the equations describing the field and therefore act to select or stabilise one manifest form from the set of forms which are possible for that type of field. (Webster & Goodwin 1996: 121)
In contrast to the Darwinist approaches discussed above, we clearly find here a distinction between the non-scientific, undefined use of the term organism (a living being) and a technical term, the form, which refers to organisms and their parts. The organism, that is the form, thereby becomes part of a scientific theory of biology, and is not reduced to genes presolved in dualisms. Obviously, Webster's and Goodwin's approach seems to be able to supply what we have been looking for: The return of the organism in the realm of biology by its representation in a biological theory! Rather than stopping here, however, we like to continue with a further question and ask: What is the status and the function of the form in that theory? Is the structuralist approach indeed the solution to developing a theory of organism and providing a foundation to biology? Or does it run into its own set of problems? Therefore, we must now take a closer look at the structuralist project and the introduction and function of form in biological theories. As we have seen, morphogenetic fields, described as mathematical representations, are the rational of form. The mathematical formula as the ideal representation of the space of possible empirical forms is related to these empirical forms as an ideal to its realisations:
In principle, therefore, a set of determinate empirical forms is conceived as being generated by a series of determinate fields and these fields are the same in the sense that they are all susceptible to description in terms of a single set of fundamental equations. In that sense they constitute a single kind. (Webster & Goodwin 1996: 121)
The generation of the single empirical forms is therefore thought of as the abstract representation of concrete embryogenetic processes. Consequently, the structuralist approach takes the complete life cycle as the object of transformation. It stands in contrast to traditional systematics,
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which is fundamental to Darwinist approaches and which focuses preferentially on adult life stages. The structuralist life cycle is represented as transformational series, and the rationale for the series are the fundamental equations which were stated above. The methodological status of these empirical forms is defined as natural kinds. The generation of the respective forms depends on the understanding of the relation between the natural kinds (as, to follow Cassirer, the form of forms) and their respective realisations. This natural kind concept, which underlies the structuralist approach, is in some respects a classic one (e. g. to the considerations of Quine 1975). To quote from Webster and Goodwin:
Consequently, the determinate members of the series are variants of that kind [the natural kind, which is given by the fundamental equations: the authors] related as transformations they comprise a rational system and the basic field equations are, in Cassirer's terminology13, a symbolic representation of the form of the series in a sense, the essence of the kind. (Webster & Goodwin 1996: 121 pp)
And furthermore:
Thus, insofar as any set of specifically different empirical forms or patterns can be understood as being produced by the same generative mechanism in this sense, that set can be conceived as comprising a single kind. From this perspective, kinds are theoretically significant kinds, that is, natural kinds. (Webster & Goodwin 1996: 121 pp)
Seemingly, the structuralist theory allowed us to cut nature at its joints. The determination of the respective natural kinds becomes possible with reference to the mathematical description. All empirical forms that follow the same basic field equation are representatives of the same kind. So, the sameness of empirical forms is defined in terms of the respective transformation rules. And this sameness implies (as was stated in the citation above) the identity of the generative mechanism. In a next step, the form becomes the realm of rational systematics. The description of forms (which, as should be remembered, refer to living entities and their parts) is used to generate a rational systematics that expresses the correct order of nature itself:
Hence, the research project of rational systematics would consist, in principle, of an attempt to reconstruct the hierarchical system of empirical taxa as a rational system in terms of the serial forms which are possible for each kind of field in the temporal sequence. (Webster & Goodwin 1996: 123)
Based on the principles of a rational systematics, a correction of traditional, empirical taxonomy is even assumed to be possible:
However, we should bear in mind that, given the dialectical relation between taxonomic and explanatory knowledge, one outcome of such a project, supposed it to be successful, might be that the merely empirical classification would be subject to theoretical corrections of a more13
One might very well wonder whether this reference to Cassirer is not misleading. However, an analysis of Cassirer's terminology in regard to its application in the structuralized approach cannot be done in the framework of this paper. To Cassires concept of the Form of Form see Gutmann & Weingarten (1996).
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or-less radical nature, as has happened in the physical sciences, since the critical (essential) properties used for distinguishing kinds will be those which are grounded by the explanatory theory. Thus, what from an empirical perspective may appear to be two distinct kinds may come to be regarded, from a theoretical perspective, as simply variants of a single kind and vice versa. (Webster & Goodwin 1996: 123)
The very subject matter of such a rational taxonomy is then the organism and its parts which is now understood as a realised form of one of the forms of forms, which were called natural kinds. In a way, this parallels the hierarchy argument, insofar as the taxonomic hierarchy is reinterpreted in terms of organismic development:
Thus, in terms of the type of morphogenetic theory proposed in this book, a qualified version of von Baer's thesis would lead us to suppose the existence of a temporal sequence of kinds of morphogenetic fields where each kind of field corresponds to a level in the empirical taxonomic hierarchy. (Webster & Goodwin 1996: 122 pp)
Here, Webster and Goodwin go a step further by proposing an existence of a temporal sequence of kinds of morphogenetic fields. In other words, they claim that the resulting rational systematics would allow the identification of evolutionary relations. In stark contrast to classical Darwinist approaches then, evolutionary development would be inferred not from genealogical relations but from the sameness of the respective transformation sets. As an example of such a procedure Goodwin uses the development of vertebrate limbs, which serve here as the units of a rational systematics:
Vierfuer-Extremitaten sind definiert als die Menge moglicher Formen, die gema den Re geln der fokalen Verdichtung, der Gabelung und der Segmentierung im morphogenetischen Feld der Extremitatenknopse erzeugt werden. Alle Formen sind aquivalent bezuglich Trans formationen, die nur diese generativen Prozesse verwenden. Damit gelangen wir zu einer logischen Definition der Vierfuer-Extremitaten, die unabhangig ist von der Stammes geschichte. (Goodwin 1997: 237)
The important point here is that the units of the rational systematics are introduced as being the same they are all vertebrate limbs in regard to a particular equivalence relation the transformations, which are used by only these generative processes for the development of the respective limbs. At this point we meet the first severe methodological problem, which becomes evident by a closer look at the procedures suggested. The procedure, which is being used here, is that of abstraction. An abstractor is a word that indicates a shift in the level of language (for further reading s. e. g. Hartmann 1993, Lorenzen 1987). If things are the same in some respect, we may introduce an abstractor referring to this particular sameness. Obviously, this does not imply that these things are identical, but only that they are the same with reference to a particular equivalence relation. Likewise, the sameness of living entities can only be stated if an appropriate language is provided. Let us look at the order of description in detail: If we describe the limbs of fishes, amphibians, reptiles and mammals as members of the same set (tetrapods), we build an abstractor referring to their sameness with respect to classical descriptive terms (e. g.
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phalanges, tarsalia, carpalia etc.). Consequently, we neglect the differences in detail, and use the same term for specified parts of the limbs of different animals. Now, in the structuralist approach sameness is not related to classical descriptive terms, but to mathematical descriptions, which were derived from descriptive terms. That is, we must start with this classical language of description, and then construct a second language level that includes the mathematical description of these parts. The next step uses the mathematical descriptions, the forms, to derive transformational relations or transformation sets of these forms. Now, according to the quote above, these transformation sets provide the equivalence relations, relative to which sameness of forms (mathematical descriptions) can be stated, and which are being used to build the rational systematics. This demonstrates that we arrive at these transformation sets and thereby at the equivalence relation by using a particular order of steps. When Webster and Goodwin state, however, that the empirical forms (i. e. the mathematical descriptions of single living entities or their parts) are the expressions of (or are derived from) the basic or fundamental equations, they reverse the methodical14 order and thereby commit a fallacy of category. The mathematical descriptions refer to living entities or their parts (e. g. the limbs of tetrapods), but the entities so described are not constructed in this way, which is described by the appropriate abstract representation. This formalist fallacy is basically grounded in a confusion of levels of descriptive language. If the difference between these levels is neglected we lose the possibility of distinguishing between empirical, formal (e. g. logic) and ideal (e. g. mathematics) sciences. Kant saw this problem with complete clarity. He insisted that biological descriptions of Naturwesen (natural entities) are of a reflexive origin:
Gleichwohl wird die teleologische Beurteilung, wenigstens problematisch, mit Recht zur Naturforschung gezogen; aber nur um sie nach der Analogie mit der Kausalitat nach Zwek ken unter Prinzipien der Beobachtung und Nachforschung zu bringen, ohne sich anzumaen, sie darnach zu erklaren. Sie gehort also zur reflektierenden, nicht der bestimmenden Urteil skraft. (Kant 1983: 306).
However, he goes on:

Es ist namlich ganz gewi, da wir die organisierten Wesen und deren innere Moglichkeit nach blo mechanischen Prinzipien der Natur nicht einmal zureichend kennen lernen, viel weniger uns erklaren konnen; und zwar so gewi, da man dreist sagen kann, es ist fur den Menschen ungereimt, auch nur einen solchen Anschlag zu fassen, oder zu hoffen, da noch etwa dereinst ein Newton aufstehen konne, der auch nur die Erzeugung eines Grashalms
14
The difference between methodological and methodical is very important for the entire argument. The word methodical means applying a method. The word methodological means thinking about methods and procedures and their application. The point of view we adopted in this article is that of a methodical culturalism. Consequently, the construction of a line of argument has to be done methodically, e. g. by starting within everyday lifeworld and obeying the principle of methodical order (for further reading s. Janich 1992 & 1997).
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nach Naturgesetzen, die keine Absicht geordnet hat, begreiflich machen werde: sondern man mu diese Einsicht dem Menschen schlechterdings absprechen (Kant 1983: 352)
This means that the description of natural entities, in terms of mechanische Prinzipien (mechanical principles), constitutes the named analogy and provides the possibility of knowledge of organised entities. But the formal description of natural entities15 has only a regulative function (in contrast to a possible constitutive function16), and therefore the construction of the formal language of description must not be confused with the natural entities themselves. Clearly, we do not have to arrive at Kant's own solution of this fundamental biological problem. However, as the above analysis of the structuralist approach has shown, we may safely assume with Kant and thus summarise the results of the analysis so far that the description of living entities or their parts in terms of a specified language (e. g. a mathematical or a physical one) is an act of attribution (s. Gutmann 1999). This attribution is composed of two distinct language levels. The formalist fallacy we have identified consists in confusing the language level of the analogy with the language level of the described entities. Obviously, the sameness of things in terms of a specified set of descriptive means does not imply the identity of these things in terms of a different set of descriptive means. So far, we have seen in our reconstruction of the structuralist approach that the introduction of the organism into biological theory is methodically flawed. The main problem was and this is an important point a confusion on the level of language! Now, we will turn to a further problem, that of a structuralist foundation for biology, which is suggested by the program of a rational systematics that could be used as corrective agent for empirical taxonomy. Such a program must supply criteria that distinguish between correct and false systematisations, so that we may avoid the following three, equally inadequate, solutions: 1. If we use the classical taxa as guidelines for our corrections, we produce a reconstructive circle which anticipates the contingently existing taxonomy as the aim of the rational reconstruction. This would be a fallacy: Since an indefinite number of descriptions of a thing (and a fortiori, of a living entity) is possible, other taxonomies may be constructed as well as the one contingently given. 2. If we neglect the classical taxonomic implications we must concede that the proposed theory of organismic transformation is the correct theory
15
Kant aims specifically at the description of natural entities in terms of mean-end-relations. But irrespective of the special description, the methodological problem is the same. 16 Kant emphasises this aspect of reflection several times: Der Begriff eines Dinges, als an sich Naturzwecks, ist also kein konstitutiver Begriff des Verstandes oder der Vernunft, kann aber doch ein regulativer Begriff fur die reflektierende Urteilskraft sein, nach einer entfernten Analogie mit unserer Kausalitat nach Zwecken uberhaupt die Nachforschung uber Gegen stande dieser Art zu leiten und uber ihren obersten Grund nachzudenken; (. . .). (Kant 1983: 324)
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of evolution. If this assumption is not justified then a dogmatic argument seems to be unavoidable17. 3. Before we can describe living entities in terms of a formal language, they must be described on a different language level that allows us to introduce the predicates needed to determine the sameness of the respective entities in terms of their evolutionary relation. The problem stated in the first inadequate solution is realised by the structuralists themselves:
If this project [the rational science of transformation: the authors] were to be successful in demonstrating that different fields could be conceived as different natural kinds and each kind corresponds to a part of the empirical taxonomic system then something like the traditional view would be vindicated. (Webster & Goodwin 1996: 123)
The criteria according to which it is possible to distinguish between those parts of the traditional empirical system that can be justified as the aim of reconstruction, and those that are rejected by the reconstruction, must indeed be given before we start to reconstruct. On the other hand this selection cannot be done by referring to the results of the structuralist approach alone, because then we run the risk of adopting the second insufficient solution: a dogmatic presupposition. If we want to avoid the accusation of a dogmatic argument, we might assume that the transformative lines that result from developmental research in terms of fundamental equations and their respective realisations are themselves already evolutionary lines. The hierarchy of developmental processes is then thought to reflect evolutionary transformations. Following the structuralist concept of evolution, the generative transformations intended here are juxtaposed to the classical historical assumptions:
When taxonomy is treated as genealogy, then the logical relationship between ontogenetic pathway and taxonomic distance is lost, because neighbouring species are then determined by historical sequences rather than by generative relationships. The historical succession of species has no obligation to proceed systematically through a space of possible forms. Though there will be a tendency for the neighbourhood relations to attractors to be expressed in the sequence of species discovered by evolution, parametric variation can be discontinuous and jump from one domain of parameter space to another, missing intervening attractors, which may be encountered later. So genealogical taxonomy a la Darwin, will in general not be the same as a rational taxonomy. (Webster & Goodwin 1996: 254 pp)
What is being suggested here is the following: the generative transformations constitute the possible morphospace, and these transformative lines are defined as evolutionary lines, which do not have to coincide with the historical succession of species. Of course such a procedure is possible, but encounters a severe methodological problem: within the line of evolutionary transformations, it is not possible to determine the later and the earlier events. That means: Webster & Goodwin offer us a solution
17
To justify an approach as the correct theory of evolution, the criticism of the concurring approaches is not sufficient. It is merely a necessary prerequisite.
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to incorporate the organism (as form) in biological theory at the cost of making evolution ahistorical18! The problem we encounter here is the problem of the Lesrichtungskriterium (criterion of evolutionary direction), which has been a core problem in evolutionary approaches. In the search for such an adequate criterion, Goodwin derives a decline in symmetries and an increase in complexity from the sequence of vertebrate limb developments. He writes:
In diesem evolutionaren Trend (the development of vertebrate limbs, the authors) zeigt sich eine allmahliche Abnahme der Symmetrie und ein stetiges Anwachsen der Komplexitat, das mit der Zunahme der Symmetriebrechungen in der morphogenetischen Kaskade einhergeht. Dies ist die naturliche Fortentwicklung eines dynamischen Systems, dessen Parameter infolge der zufalligen Neukombination des Erbguts verandert werden. (Goodwin 1997: 240)
However, this general trend of evolution, which Goodwin uses to reconstruct the correct sequence of generative transformations (i. e. the sequence of the correct ontogenetic transformations), must be justified again. For example, one might very well argue against the criterion of increasing complexity on empirical grounds: clearly, we know many cases of so-called reduction lines, e. g. such curious forms as hemichordates, pogonophores or sipunculids. But even if we leave these empirical arguments aside, the criterion (increase of complexity) is still problematic. It has the methodological function of a premise, i. e. it must be known before we start to reconstruct. Let us consider that the criterion itself is (at least) a three-termed comparative (polar contrary to simple) predicate. If we consider a thing A to be complex, we actually mean: A is more complex than B with regard to a criterion C, which serves as the relevant parameter for comparison. If evolution is defined as the increase in complexity, the comparability of the compared things (here the living entities) must be guaranteed, and the criterion of comparison determined beforehand. If we use knowledge gained from the classical taxonomic concepts in order to determine e. g. the route from specific fishes as predecessors to specific amphibian and reptile forms, then the well-known problem of a vicious circle arises (see Ax 1984). Look for example at the following quote from Goodwin:
Wir konnen einen dieser Trends in der Evolution der Anhangsel von Wirbeltieren identifizie ren. Offensichtlich entwickeln sich die Fischflossen aus seitlichen Hautfalten, die sich an beiden Korperseiten der fruhen Wirbeltiere entlangzogen. Diese Falten bildeten sich dann zu zwei paarigen Anhangseln zuruck, den Brust- und den Bauchflossen. Bei den Knochen fischen, zu denen auch die Quastenflosser gehoren, schrumpften diese weiter zu gelappten Flossen, welche am Anfang der fur die Vierfuer, die sich aus den Quastenfloern entwickel ten, kennzeichnenden Gabelungen stehen. (Goodwin 1997: 240)
The increase in complexity in this case refers to a particular line of evolution, namely from early fishes via e. g. coelacanths towards early tetra18
This shortcoming could be avoided if a biogenetic principle is stated. In this case, a vicious circle will appear when the principle itself has to be derived or justified.
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pods. Despite the numerous empirical problems that are connected with this reconstruction (particularly whether coelacanths are the predecessors of modern tetrapods at all), the evolutionary line has to be justified as a valid empirical assumption. In order to justify such a premise, we have to refer to knowledge, which is based upon the classical languages of description. Therefore, if we defined evolution as a process of increasing complexity this way, we would commit a petitio principii, which is as unconvincing as the alternative options. At the same time the function of the structuralist approach, namely to serve as a corrective to classical empirical taxonomy, gets lost. The fact that the structuralist description of living entities in terms of a formal (here mathematical) language refers to classical implications for the direction of evolution which is, in a second step, described as an increase in complexity leads to the third inadequate solution we indicated above. As we stated, an adoption of the classical determinations of the direction of transformation (in our example the line from early fishes via coelacanths to early vertebrates19) forces us also to adopt the underlying language of description. This difficulty is seen within the concept as the difference between naming and interpretation. Thus, Webster and Goodwin give an example of a classical concept, the names of structures (such as phalanges or digits or ulna):
The original tetrapod limb, that which belonged to the common ancestor of the fourlegged lineage, is assumed to have been one with five digits, since this is the condition of primitive, extinct amphibians such as Eryops and continues to be the predominant amphibian limb form. These digits were accordingly named 1, 2, 3, 4 and 5. Darwinian belief that there is some type of inherited, material continuity of the individual elements that constitute a persisting form, such as the tetrapod limb, results in the view that the elements in the limbs of different species can be named in relation to the original digits. (Webster & Goodwin 1996: 140)
Here, structures are thought to be homologous referring to an original type which is genealogically connected to them. The original type serves as an archetype here, which allows us to determine whether or not a structure possessed by two animals is homologous (for the archetype problem s. Young 1993, Gutmann 1996). Because the type is thought to exist (here in the form of Eryops as its representative), the respective structures (here the phalanges) are equally thought to exist as a pattern. This point of view provokes a lot of methodological difficulties that have previously been identified and discussed (Weingarten 1992, 1993). The structuralists contrast this position with their concept of homology as an equivalence relation:
The invariant is simply connectedness. The mappings can obey the constraint that the angles between intersecting lines in one space are preserved in the image space. These define equiva19
In case of this forms there may even be a minimum consensus. This is surely not the case regarding the wide filed of invertebrates.
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lence under conformal mappings. Or distance can be preserved, as in the motions of rigid bodies (translations, rotations and inversions), giving equivalence under metric transformations. Each equivalence set has an invariant property that is preserved by the transformation. A hierarchy of such equivalence classes can be defined, each successive invariant including but imposing more constraints than the previous. (Webster & Goodwin 1996: 144)
If the homology is introduced this way, the structuralists avoid the obvious methodological shortcoming of the classical concept. But on the other hand they introduce the equivalence relations with reference to descriptions of living entities, which already contain such terms as digits, phalanges and ulna. These terms indicate the evolutionary comparability of the structures. So, in a way the comparability of the natural entities being described is already stated, by describing them in classical terms. What is missing in the structuralist approach is the construction of a reference language that allows us to operationalise our formal description. Metaphorically speaking, the entry point of the structuralist solutions to the naming problem is too high. Consequently, the biological meaning of structures that are described as homologous this way cannot be derived from the formal description. At this point we can summarise the results of our reconstructions of the examples of contradictory and mutually exclusive points of view, namely the Darwinist on the one hand and the structuralist on the other. They have led us to a curious position. When we regain the organism as an intelligible biological unit in terms of structuralism history collapses into mathematical transformations, whereas the attempt to argue from a genealogical or historical point of view as performed by Darwinist approaches results in the disappearance of the organism. At the same time, our reconstruction showed that neither result is the product of the nature of nature or the nature of organisms, but are tightly connected with the language we use to describe the respective assumed realms of biology (life on the one hand, the organism on the other). The result, that the language we naturally apply becomes a part of the problem that it was supposed to solve, provides us with a new starting point for the construction of a theory of organism and the foundation of biology.
4 Looking for a New Starting Point

So far, we have analysed different approaches in terms of their efforts to introduce a scientific concept of organism into biological theories. We have shown that none of the suggestions was completely satisfying. Ultimately, the language applied was itself part of the problem. Since we want to suggest a new approach to conceptualising the organism and make it accessible to biological theorising the obvious starting point for such a project is an analysis of the language used. We need to begin with the very first question that was the reason for these considerations: What is an organism?
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The form of this question is typical of What is . . . questions, e. g. What is a chair?, or What is a cow? The simple form of these questions indicates that the questions refer to concrete things. Usually, such questions would be answered by enumerating some characteristic features. In the case of the question What is an organism?, we might give a list of several important or relevant features. Those features include several abilities such as metabolising or reproducing. One argument against this approach is that by using these features we gain nothing more than a synonym for living entities. A more serious argument is that the descriptive terms metabolise and reproduce come from an already highly standardised scientific language in contrast to the word chair or the feature used to sit upon. Adding here the ability to have a form as proposed by the structuralists would not make the situation any better. The next approach to this problem must therefore be to have a closer look at the second part of the expression living entity, namely the word entity. It was used exclusively as a substitute for such expressions as plant or animal, and it is exactly this substitution that leads us to the starting point we need. In contrast to life or entity which could be understood as standing tor specific ontological or even metaphysical intuitions the terms animal and plant are tightly connected with the everyday lifeworld. The term lifeworld, with its qualifier everyday, has been used several times throughout this paper, and its meaning was probably understood intuitively. Now, however, is the time to introduce the term more explicitly. Everyday lifeworld refers to the practices that comprise a wide field of differing human activities. The means and tools that are used within these fields of activities as well as the respective results constitute an array of mean-end-relationships. It is important to note here that this field of human activities and their means-end-relationships include language as well. Consequently, the term everyday lifeworld describes such different activities and techniques as the building of houses, weaving, mining, metallurgy, the cultivation of weed and the breeding of cattle. Obviously within different specific contexts of the everyday lifeworld, we can successfully deal with building material, yarn, ore, animals and plants and the procedure of naming them, i. e. we are able to achieve our goals and ends. This might look like a trivial truism, but it is of some methodological relevance. It is exactly at this point that we observe a significant asymmetry in the relation between science and the everyday lifeworld. In the contexts of everyday lifeworld we are clearly able to deal with animals and plants without having to refer to scientific (here biological) knowledge20. A scientific treatment of animals and plants, however, is impossible without reference to everyday knowledge and know-how.
20
This does not exclude the utilisation of scientific knowledge at all. As we will see soon, e. g. physical (and chemical) knowledge can be used, insofar as the constitution of these forms of knowledge is not based in relevant aspects on biology.
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One result of this asymmetry between the everyday lifeworld and science is that the everyday lifeworld precedes science methodologically, and the language of the everyday lifeworld precedes scientific language. Within the everyday lifeworld we usually use a language which permits communication about the relevant aspects of living entities with which we are usually confronted. What is even more important is the fact that we can successfully handle, manipulate, breed, cultivate etc. living entities and describe the relevant operations, within evervday language. The methodological priority of everyday language is reflected by the fact that even if we want to answer scientific questions, we will necessarily start with those parts of everyday language that belong to this field of successful cultural practices. Since we diagnosed the theoretical problem analysed in the preceding chapters as largely due to difficulties with language, we can now see that following a methodological order of everyday lifeworld language and scientific language can prevent such language difficulties. In other words: starting from everyday lifeworld language (which is rooted in cultural practices) we can develop a procedure that allows the distinction of at least two levels of language21, which are ordered insofar as everyday language and practices provide the methodical starting point for the introduction of the primary terms into scientific language. The language of description that is designated as a scientific language here refers to particles of the everyday lifeworld language. These particles originate in the practical context within which we deal with living entities. When we deal with living entities on this level of the everyday lifeworld, the term living entities encompasses simply the animals and plants with which we are usually acquainted particularly within such explicitly marked practices like breeding and cultivation. Referring to the relevant particles of language which shall be abbreviated as L (Lifeworld language particles) we can describe properties (for example features, motion, performances or behaviours) of animals and plants that are relevant for different purposes. These purposes reflect the respective goals we aim at within different practices. An example might illustrate this point. The ability to carry burdens, possessed by the group of cattle-like animals, is of the utmost importance if one's aim is to produce (breed) animals with optimal performance in this area. In other circumstances other purposes are chosen, for example, the quantity of milk that can be produced by cows, the productivity of plants, the beauty of the fins of goldfish. In all these cases we are able to describe the relevant features (milk production, fin beauty) of these animals by referring to the goals we attempt to achieve. Usually these goals can be identified as the direct or indirect purposes of breeding or cultivation, and it is pre21
Obviously there are at least two levels of language; we needn't stop with the construction of further language levels at this point. The respective higher language levels have to be considered to refer to the respective lower levels. The most important point again is the startingpoint named as a methodical starting point or beginning.
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cisely these practices, which have accompanied the historical development of human societies, that give rise to the relevant knowledge and knowhow. In the same way, breeding and cultivation practices provide the necessary language particles that we need to describe animals or plants. These language particles refer not only to individual properties, like shape or physical abilities, but also to super-individual properties, like the ability to propagate. All these properties, features, abilities etc. are then described by reference to the respective manipulative action that constitutes breeding or cultivation. Consequently these descriptive terms refer to purposeful human action and not to natural aspects of the respective living entities. Obviously the language refers to specific operations (usually within the context of use and production of animals and plants). The relevant particles of this language are introduced not just by means of description but by reference to the relevant practices. We can call this a prescriptive point of view in contrast to a pure descriptive one. Note that what we called the features, characters or behaviour of animals and plants are descriptions (i. g. prescriptions, see above) within the marked context, which do not imply biological knowledge. Consequently, the knowledge and know-how we gain within the context of the everyday lifeworld has a restricted claim of validity in several important respects. In stark contrast to the field of scientific statements and predictions, we neither expect nor demand universality but are restricted to particular validity. In the next section we will show how a scientific language a language of description can be acquired by starting from the language of the everyday lifeworld.
4.1
Towards a Language of Description
To obtain adequate scientific explanations, descriptions or predictions, we have to introduce a standardised language language S that both preserves the knowledge and know-how of the primary language level L and also avoids all the indeterminacy, ambiguity and sometimes even inconsistency, of this language L. Consequently, the standardised language we are looking for must be constructed by starting within the restricted context of the everyday life world, and transcending it by explicating a procedure that provides a sufficient degree of reproducibility and clarity. To accomplish such a task we suggest applying the constructive model theory (for further reading see Gutmann 1996, 2000). In this approach, a model is not merely a two-termed relation that constitutes an analogy, e. g. between technical abstractions and several properties (for example the shape or performance of living entities), as is suggested by such advanced approaches as the structuralist theory of organism. A model in terms of the methodical culturalist theory is a three-termed relation, as the following example shows:
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As we know from everyday lifeworld experience a horse can be used to pull a plough. It is possible to describe this action in terms of classical mechanics. We then describe the horse as if it actually were a force-generating unit. We do not simply identify horses with machines or engines (in terms of force-generating units) but we describe them as if they were machines, i. e. we use machines to introduce such terms we need to describe the motion, action and performance of the horse in mechanical terms. More generally, using the three-term relationships in methodical culturalist model theory, an object whose relevant aspects are adequately described within one language, is described by applying particles borrowed from another language. This modelling will result in the construction of a (in this case mechanically) standardised language S, by applying a technical language. By modelling a horse as a force-generating unit we ascribe functions to several relevant parts of the animal. Because we introduced a model as a three-termed relation, we can name the parts of the animal that are relevant to a specified motion, action or performance (described in terms of L where L stands for everyday lifeworld language) as structures that perform a specified function (described in terms of S where S stands for standardised, e. g. scientific, language). That is: the application of the technical knowledge and know-how by means of language denotes a procedure that can be called structuralising. In applying the procedure of methodical culturalist model theory, we move from one language level (L) to another (S). This shift of language level is marked by the shift of language particles. Whereas in L we deal with animals, plants, living entities and their component parts (such as their limbs), in S we deal with their structures and organs (for the introduction of the term organ see Gutmann 1996). This procedure can be applied to the other descriptive terms we mentioned above (such as motion, movement and performances of living entities) if we only keep in mind that the everyday lifeworld contexts from where we start are themselves practical contexts. That means that living entities neither have parts, nor do they have structures and organs. Rather, we describe the objects in L (living entities) on the one hand, functions, structures and organs we intend within the specified context of inquiry in S on the other hand. Having found the place of the word living entities in everyday lifeword language, these considerations allow us now to determine the methodical status of the organism as well.
4.2
The Term Organism
Based on the preceding considerations, we now suggest the introduction of the term organism into biology as a concept of notion or Reflexionsterminus (for further reading s. Gutmann 1996, Janich & Weingarten 1999, Gutmann 2000). Referring to Kant's explication of the description of Naturwesen (natural beings) in terms of means and ends, we can
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describe living entities (starting with the well known plants and animals of everyday life) as organisms. In doing so, the organism is constructed by beginning with the single or collective properties or characteristics of living entities (animals, plants etc.). These properties or characteristics are used to constitute primary biological terms, which we call organismic properties. With reference to organismic properties we suggest using the term organism as an abbreviation to stand for a list of all those properties or characteristics of living entities described as organismic properties. Using the term organism in such a way, it becomes part of a meta-language M. M refers to Sb (where b stands for biological), and describes the constituted primary and subsequent constructed biological terms, which in turn have their foundation in everyday life practices in L. This sum of organismic properties should be seen as an open list of the organismic properties, which can be expanded if necessary (e. g. for new biological disciplines). In a certain sense, the notion of organism is replaced by organismic properties. Organismic properties can be defined operationally, e. g. by the model-based constitution of the universe of discourse of specified biological theories, beginning with the qualities or movement of animals and plants. Such a methodological reconstruction of biology leads to distinct and well-defined branches like functional morphology, constructional morphology, reconstructional morphology, constructional genetics, constructional theory of evolution etc. The functional and constructional description of animals determines an organismic construction. The same is possible for collective properties like the ability to propagate. In the next section we will try to make this general description of the methodological reconstruction of biology clearer by using an example taken from morphology.
4.3 Presenting a Case Study: How does a Crayfish get its Structures?
To give an example of what might be called an organismic description of an animal, let us take a closer look to the procedure of structuralisation. Within everyday lifeworld contexts we are well acquainted with interesting animals such as crayfish22. These animals are interesting for several reasons. They are of great economic interest, they can easily be farmed, and last not least, when carefully prepared many people find them very tasty. Crayfish early became a preferred object of biological description and research, particularly in neurophysiology, because they show characteristic escape behaviour (s. e. g. Kennedy & Takeda 1965, Wine & Hagiwara 1977, Wine & Krasne 1982). This behaviour, which is also observed in
22
The following from Gutmann (1998).
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Fig. 1. Schematic representation of some anatomical aspects of the thorax of a crayfish (P. leniusculus). Longitudinal section. (After Gutmann 1998)
other crustaceans such as prawns, consists of the abrupt flexion of the bottom of the crayfish's tail against the bottom of its forepart. If we want to know how this motion which usually results in locomotion occurs, we could start by describing this process as propulsion. However, there is a great difference between a body that is accelerated by the application of an external force (like a stone thrown or catapulted) and a crayfish that moves by itself. The question here is: how are these forces produced from within the crayfish? We can approach this question by identifying the crayfish as a force-generating unit. We then assume that parts in the unit crayfish act in concert with each other to generate the force (propulsion). We assume that the crayfish can be structuralised as a force-generating unit. To test this assumption, three steps are necessary to enable us to structure the crayfish in the required form. First, we have to section it, in order to identify anatomically the parts of these animals. Second, we must provide a model that allows possible functions to be ascribed. Finally, we have to integrate the resulting structures within a mechanical framework that is termed the construction of the animal. Before we go through the suggested procedure in detail, it is important to clarify what can be expected from the scientific investigation of the crayfish's escape behaviour. At the end of this procedure, we should not assume that the crayfish is a force-generating unit, or that a crayfish is built up out of the force-generating structures identified. In this case we would simply but wrongly try to create a synthesis out of our analytical
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functional ascription and structuralisation; an analytical procedure we chose (from alternatives) in order to explain particular aspects of the behaviour23 of interest. Such a reversion from analysis to synthesis is wrong because it results in a fallacy of category. It assumes that the synthesis (something is made up of y) is the inverse operation to analysis (something can be partitioned into y). To avoid such a fallacy, therefore, we will have to keep in mind that sectioning is itself a practice. Within this practice we do not use descriptive terms alone. In contrast, we are elaborating the parts of the animal and particularly its inner parts by applying an experimental procedure. During preparation we follow the extension of tendons and muscle loops with a probe, we examine the stiffness of cartilage or skeletal parts by bending and pulling, the strength of muscular and connective tissue attachments, etc. Finally, we do not describe the animals per se, but the results of our more or less successful operations and actions. Even considering only this simple procedure of sectioning (and not mentioning the advanced techniques of modern histology, etc.), we can state that morphology is not simply a descriptive science. With these cautionary remarks we can now have a closer look at the steps of structuralising, and we begin with the first step: sectioning. 4.3.1 The Thoraco-Abdominal Framework The crayfish24 body can be subdivided in two parts, the cephalothorax and the abdomen. The cephalothorax bears the antenna, the mouthparts and the pereiopods (i. e. the thoracic walking legs). The carapace connects the apical head capsule with the anterior aspects of the first abdominal segment. The thorax is equipped with a well-developed endophragmal system (see Fig. 1). The apodemes of the endophragm connect the sternum with the epimeral wall. In both cases, the endophragm provides attachments for
23
Behaviour is used here as an everyday language expression, and not as an ethological terminus technicus. 24 Crayfishes (Astacida) are members of a group of decapoda, so-called macrura, because of their well developed tail (the abdomen). As representatives of the higher crustaceans (malacostraca), they may be of some importance for the systematic understanding of the entire group (for further reading Albrecht 1982, Scholz & Richter 1995, Schram 1983, Gruner 1993, Dahl 1983). Additionally, Astacida could be relevant for the identification of possible brachyura-origin within the macrurans. The decapoda were the subject matter of anatomical and morphological description very early in the history of biology (s. Milne-Edwards 1851). For the sake of the argument, we developed above, we will give a short sketch of some main anatomical aspects of macruran anatomy, referring to the standard literature and own results (Huxley 1881, Netz 1917, Schmidt 1915, McLaughlin 1977, Holdich & Reeve 1988; for other macrurans s. Daniel 19291932, Felgenbauer & Abele 1983; for other arthropods s. Manton 1977, Boudreaux 1979). Because the underlying inquiries refer to different representatives of Astacida (e. g. Astacus astacus, Astacus leptodacytlus, Austropotambius torrentium, Procambarus clarkii etc.), we will only talk about crayfish here abstracting from the differences in detail that can be found between each of these forms. For the examination of the musculature of P. leniusculus that was the standard form here, longitudinal as well as cross sections were necessary (for further reading s. Gutmann 1995).
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Fig. 2. Schematic representation of some anatomical aspects of the abdomen of a crayfish (P. leniusculus). Longitudinal section. Only some aspects of the superficial and the pleopodmusculature are considered. (After Gutmann 1998)
the pereiopod musculature. The dorsal parts of the endophragm act as attachments for the oblique musculature, too. The thoraco-abdominal hinge provides the connection between thorax and abdomen (Baumeister 1985, Netz 1917). The entire thorax is thus very rigidly structured and provides a stiff framework for the insertion of pereiopods as well as dorsal and ventral abdominal musculature. The abdominal skeleton is composed of a series of six very rigid, cuticularised straps, each of them consisting of an (upper) tergital and a (lower) sternal part. Each segment is connected to its following segment by flexible intersegmental membranes (see Fig. 2). 4.3.2 The Abdominal Musculature The crayfish tail is more or less dorso-ventrally flattened. This shape is described25 as compressed. When we examine the crayfish tail in longitudinal section, we find a mass of entwined muscles, running predominantly from the thorax and the carapace to the last segment and the telson. The muscles are tightly attached to the sternal and tergital aspects of the skeleton (see Fig. 3). We can identify two main portions of muscles: the dorsal and the ventral musculature. The dorsal musculature consists of two series of paired and entwined muscles on each side of the abdomen connecting each tergital plate with the next, starting from the medio-lateral aspects of the carapace and the epimeral wall respectively. The dorsal muscular ropes begin within the dorso-medial parts of the thorax and end on the inner dorsal aspects of the tergit of each segment. In contrast to the dorsal musculature, which can be described in rather simple terms, the ventral musculature is at first glance a very confusing arrangement of muscular fibres running in different directions. Anatomi-
25
Shape is a term in L in this context, because we do not deal with the form.
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Fig. 3. Schematic representation of the main anatomical aspects of the abdominal musculature of a crayfish (P. leniusculus). Longitudinal section. See text for further explanation. (After Gutmann 1998)
cally, we can identify four main aspects, namely the oblique, the transverse, the dorso-lateral and the central muscles (see Fig. 3, 5). The transverse muscles run from one side of each segment to the other, connecting both sides near the respective hinge points. The central muscles are paired and entwined, running from each segment to the next, connecting the transverse muscles. The so-called transverse membranes are integrated within this framework of longitudinal and transverse musculature. This system spans the median aspects of the crayfish abdomen. It connects the lateral aspects of the tergits near by the hinge points with the lateral aspects of the respective central as well as dorso-lateral muscles (Rayner & Wiersma 1967, Pilgrim & Wiersma 1963, Schmidt 1915). The transverse membranes provide important attachment surfaces for the predominant form of musculature that can be found in the crayfish abdomen, namely the oblique muscles. The massive oblique muscles run rostro-caudally, each spanning two segments or hinge points (except for the last two muscles; see Fig. 5). These muscles connect the posterior aspects of ventral thorax (above the endophragm and the lower parts of the epimeral wall) with the sternal skeletal straps (which provide the attachments for the pleopod musculature) and finally the terminal dorsal aspects of the telson. These muscles are tightly connected to the framework that consists of the transverse and the central muscles on the one hand, and the transverse membranes on the other. Their connection with the anterio-ventral aspects of the central muscles should be particularly noted, because the oblique muscles follow the central muscle loops in the opposite direction of rotation.
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4.3.3 The Procedure of Functional Ascription This short outline of some major aspects of crayfish anatomy may be sufficient to indicate its startling complexity (for further detail s. Schmidt 1915). These lengthy descriptions were directed towards the anatomical sectioning and description of crayfish anatomy. This particularly the abdominal anatomy has a rather long tradition within classical biology. We have not treated the functional interpretation of these complex muscular relations, however, and the functional interpretation has in fact been quite problematic for classical biology (see Rayner & Wiersma 1967, Rayner 1965, Balss 1940, Marshall & Williams 1972, Freuer & Graham 1976) since more or less all the muscular groups are directly or indirectly connected with each other. In order to approach our question how is the crayfish escape behaviour performed? we cannot stop at the descriptions of diverse muscles, but want to know how they are orchestrated, how they function together to produce the propulsion in question. Such a move in explanatory goals requires a shift in language. So far, we have used a purely anatomical description that allows the words muscle, membrane or hinge to be used in a terminologically loose sense. Now we need to construct a language that allows functional ascription. Applying the culturalist concept of modelling we can provide such a language of description. Accordingly, we re-describe the anatomically identified parts of the crayfish abdomen (and some of the thoracic aspects) in terms of the biomechanical language of description. The skeletal framework can be considered as a series of tube-like sections that are mechanically characterised by their stiffness. The motion of each segment relative to the next is a case of rotation around an axis that connects the hinges of both sides of each segment. In order to describe muscles as functional units in terms of force-generating units (Kraftaggregate26), and referring to the mode of force generation as tension-fibres-force-generating structures (TFGS27) it is necessary to consider the relevant parameters. These are the at least two points of insertion, the axis of rotation and the position of the TFGS relative to the insertions and axis of rotation. For the functional analysis of the tail flick, the movement or locomotion with which we started our description, we can discern two possible types of motion. On the one hand, there is the rapid movement of the abdominal ventral aspect towards the thorax's ventral aspect (i. e. the flexion). On the other hand, there is the relaxation of the abdomen to its original position (i. e. the extension). When we structuralise the anatomically described aspects of the crayfish's abdomen according to the theoretical assumptions, which we gained from the model, we can discern at least four possible functional arrangements of TFGS for each of these perfor26 27
For the methodological foundation of this ascription s. Gutmann 2000. For the Introduction of this term s. Gutmann 2000.
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Fig. 4. Four cases of flexor function. Case a represents the direct and case bd the indirect mode. See text for further explanation. (After Gutmann 1995 & 1998)
mances. According to the considerations of Rayner and Wiersma (1967), a model can be drawn that represents the four possible arrangements (Gutmann 1995 & 1998). These four arrangements (see Figure 4) represent two different modes of the same performance, which we shall call the direct and the indirect effect of the TFGSs. Case a concerns a direct acting TFGS, because both insertion points are below the axis of rotation and directly connected by the TFGSs. Case bd represent differently organised indirect TFGS actions, because in contrast to a the insertion points are either (1) not both below the axis of rotation, or (2) they are not directly connected by the TFGS, or (3) they both are below the axis of rotation and they are not directly connected. The same differentiation as in the case of flexion can be done for the extension. If we call a TGFS an agonist, referring to one of both possible performances, then we can call a TFGS that reproduces the original situation the antagonist of the first TFGS. Based upon these differentiations, we can ascribe respective functions to the muscles we described anatomically above. Accordingly, the oblique muscles can be described as direct acting flexors. The central muscles that build the second part of those TFGSs running more or less longitudinally along the anterior-posterior
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axis, are indirectly acting flexors. The function of the dorso-lateral as well as the transverse muscles remains unclear until we recognise that the contraction of these TFGSs will lift the upper parts of the oblique as well as the central muscles dorsally, because they shift the transverse membranes dorsally during contraction. The importance of their action in terms of the tail-flick mechanism appears to lie in its effect on the changes in length of the direct oblique muscles, which may increase the efficiency of the latter TFGSs (see Rayner & Wiersma 1967). Consequently both muscles can be described as indirectly acting TFGSs with neither flexor nor extensor function in a narrow sense. 4.3.4 Considerations of Constructional Morphology In summary, functional ascription allows us to identify possible functions of the TFGSs. For this procedure of structuralising we dealt with two language levels, the everyday lifeworld and anatomical language, and the functional description of the parts of the respective animal. In order to integrate all the single ascriptions, we now reach a constructional morphological description of the animal. Thereby, the integration of the functional ascription cannot be reduced to a simple addition of the single elements we have already elaborated functionally. Rather, we need a rule that governs the constructional description itself. Such a rule can be found in the norm of coherence. Since it is our effort to model the living entity as machine, the norm of coherence must be found in the rules for the correct construction of classical engines. Here we find that the construction and working of an engine requires e. g. force closure in addition to form closure and material closure28. Force closure is a term derived from the language referring to the model, and is applied to the animals or parts characterised in L as well. The particles coherence and construction belong to language S. In other words: If we deal with living entities as if they were engines (in terms of classical machines), then the coherence is the norm we need to built a correct construction of the corresponding living entity (for further reading see Gutmann 2000). The results of such a constructional description can be seen in Fig. 5 from a dorsal point of view. In order to built a coherent construction we have to remember that we started with a particular performance of the crayfish, in this case of locomotion, namely the tail-flick29. In this methodical structuralisation the starting point is reflected in characterising the construction as a bionomal construction. Referring to the constructional representation in Fig. 5, we can assume that the single ascriptions we made previously were only analytical descrip28
See Schilling et al. (1993) for the differing methodological status of these precepts or norms in the original context of Maschinenbau. 29 If we were interested in its walking we were able to reconstruct the conditions of this motion too. We concentrated here on the tail-flick alone (for further consideration s. Gutmann 1998).
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Fig. 5. The functional arrangement of the main abdominal muscles of a crayfish in a dorsoventral view. (After Gutmann 1998)
tions of the working conditions of each single muscle. For the constructional representation, we will have to take into consideration all the single structures, the TFGSs as well as the force-transmitting structures (in this case the transverse membranes as well as the skeletal elements). Consequently the construction of an animal is to be understood as a functional and a structural unit. Keeping this in mind underlines that it is not just most of the biomechanically described sets of abdominal muscles that are of indirect effect, but all of them. Even the oblique muscles will only act as flexors if the transverse membranes are directly or indirectly connected to the skeleton via dorso-lateral muscles. Additionally, the framework of central and transverse muscles preserves the working conditions for the direct action of the oblique muscles. From this point of view, the system of transverse and central muscles is not simply a frame into which the other muscles are integrated. Rather, they enable the oblique muscles to act directly as flexors insofar as they act as indirect muscles. What is true on the level of the combined muscle action can also be stated for the entire functional unit of the abdomen. If we understand the abdomen as a working unit, able to act as a propulsor in the tail flick, the working condition for this performance is provided by the tight mechanical connection with the thorax. The endophragm is built as a double-barrelled structure (integrating the ventral aspect of sternum with the dorsal epimeral aspects; see Fig. 1) that provides a rigid frame for the insertion of the massive abdominal musculature. In addition, torsion of
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the thorax is avoided a function that is executed in the abdomen by the combination of cross-section stabilising sternal straps and the lateral row of pleura at each segment. Together with the carapace, which provides a similar biomechanical structure for the lateral and dorsal aspects of the abdominal musculature, the thoracic unit is the main anterior insertion point of the abdominal musculature. The second insertion point is provided by the posterior attachment of musculature at the telson. The ventral position of this arrangement in relation to the hinge-point line is constituted by the sternal attachments on the one hand, and the integration of the main flexors into the frame of central and transverse muscle on the other. From this constructional morphological point of view, the tail-flick demands a mechanically highly integrated working unit that must be described in terms of coherence and the bionomic action of the entire construction. The construction itself was introduced within a model-guided bottom-up approach, reflecting on the working conditions at each analytical level of description.
5 Biology as an Organismic Discipline

The case study showed that we need a language that is methodically built and strictly ordered. To answer a rather simple question about the powergenerating structures of a tail flick, it was necessary to differentiate between two language levels30: 1. Everyday lifeword language, technical and anatomical description. a) At this primary language level the methodical starting point was found within the description of some relevant performances, features, motion, locomotion, and the shape of the animals themselves. The locomotion and its characteristics can be described phenomenologically, i. e. without the strict criteria we usually demand for scientific descriptions. This level permitted a nonbiological description of the living entities and their motion. b) The anatomical description used well-known terms like muscle and tendons. Because the descriptive particles of this language are introduced by applying type-standards31, we can produce language parti30
One could even speak of a striated language. We did not intend to split up each single level into several distinct language levels or sublevels because the borderlines between the functional and the constructional level that served as an example here are rather fluid. Nevertheless, one should keep in mind, that we identified a constitutive asymmetry between the functional and the constructional structuralisation, because we already need functional terms in order to introduce notions of constructional morphology but not necessarily the other way round (s. Gutmann 1996). It is of the utmost importance here, to emphasise, that the differentiation we proposed within the language levels (L or S) must not be misunderstood as ontological representations. The construction of a scientific language provides a tool for specified ends and does not represent aspects of the world (e. g. Dewey 1958). 31 For the difference between type standard and functional standard s. Gutmann (1998).
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cles that belong to the level L and not to biological theories. From this point of view, anatomy itself irrespective of the fact that physical terms are sometimes used to describe parts of animals is more an art than a science. As a result we acquire a description of the animals and their parts in terms of non-biological theories. 2. Standardised scientific languages referring to methodically ordered, regulated and standardised (e. g. experimental) practices denoted as Sb in the text. More general: a) Sb Particles of standardised language applied within biology, denoted by Sb in the text. Sbm The functional modelling and the functional and constructional description that are applied in morphology, denoted by Sbm. a) The motion in which we were interested was described by applying particles of technical and physical language. Both of these are free from any constitutive biological knowledge or know-how. b) The application of the procedure of the constructional model allowed functional ascription of the animal parts we dealt with on level 1 b. Here, we make the transition from the type-oriented anatomical to a biomechanical structuralisation in a narrower sense, using terms like force-generating units, forcetransmitting units, axis of rotation and so on. This procedure of functional ascription allows the animal to be structuralised in terms of functional morphology. c) Finally, a constructional morphological description was performed. Here we used norms of construction like coherence to obtain the constructions of the respective animals. By referring to constructions, we can even speak of the form of the organism. Sbx Further biological descriptions, e. g. of a physiological, genetical, ecological origin. b) Sx Particles of further standardised languages applied within other, non-biological sciences (e. g. physics or chemistry, which might be denoted by Sp, Sc etc.). The result of this differentiated procedure was an organismic description of a living entity in this case, a crayfish. Beyond the narrow limits of morphological interest of knowledge, many other kinds of structuralisation of living entities are possible. If we chose different performances, modes of motions and locomotion, other properties of living entities within the methodical starting point of the everyday lifeworld or if we intend to achieve different explanatory goals, we can constitute the primary ob-
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jects of other biological disciplines such as physiology, genetics, systematics, taxonomy or even ecology. These foundations, which we call protobiological, of several biological branches have previously been elaborated (Gutmann 1996). In all these cases, we construct standardised languages (Sbx) which refer to these differing types of knowledge and know-how within the relevant methodical starting point (described in L). From this point of view biology necessarily becomes an organismic science. The seemingly ontological unity of the organism is methodologically resolved in an (empirically) unrestricted number of organismic structuralisations of living entities. This highlights one of the most surprising results of our introduction of the term organism: the fact that we have acquired several distinct languages of description and structuralisation. At first glance this differentiation might be seen as a methodological separation of biological theories. One might assume that by avoiding the methodological shortcoming of the ontological considerations identified above we produced not just one single gap, e. g. between the ecological and the evolutionary realm, but an indefinite number of such gaps between each biological discipline. But in contrast, our methodological differentiation enables us to transcend the limits of each separately constituted research field. To see this, we must keep in mind that our case study showed only one single line of constitution, i. e. Sb was explicated only in terms of morphology (Sbm). The different languages summarised in Sb and expressed by the relevant index do not represent properties of living entities but are a set of tools we use to answer scientific questions. Consequently, the construction of relationships between these structuralisations is possible. In this case we construct a relationship between two languages in S by referring to a third language, that allows us to explicate our interest or knowledge, the explananda, the means we need to achieve our explanatory goals. The most important restriction on such a triplet of languages is the methodical order we have to obey, as our case study showed. From a culturalist point of view, we can reject the interpretation of methodological separationism if we keep in mind that the common starting point of all the single foundations leading to the constitution of biological objects is the everyday lifeworld itself. Unless we doubt the unity of this field of common experience even nonscientific experience the unity of biology is not endangered. This unity has to be maintained in the first step of constitution and the mode of reflection, but not in the night of abstraction. With Goethe: Wer will was Lebendigs erkennen und beschreiben, sucht erst den Geist heraus zu treiben, Dann hat er die Teile in seiner Hand, Fehlt leider! Nur das geistige Band.
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Acknowledgement
We would like to thank Jackie Leach Scully from the University of Basel, who proofread the manuscript and gave us some important hints for the improvement of our article. Special thanks goes to Christoph RehmannSutter who made possible the financial support for the proof-reading of the manuscript. The support was granted by the Stiftung Mensch Gesellschaft Umwelt (MGU) at the University of Basel, grant F 42/95.
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The Theory of Organism

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The Theory of Organism and the Culturalist Foundation of Biology

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

2 The Organism in Modern Biology Its Loss and its Defence

M. Gutmann and E. M. Neumann-Held

Organism Lost in Life

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

Organism Lost in Hierarchies The Schism of Biology

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The Status of Hierarchies for the Foundation of Biology

The theory of organism and the culturalist foundation of biology

As to the status of postulate 1.1, they state:

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

3 Introducing the Organism as a Unit of Transformation

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

However, he goes on:

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

4 Looking for a New Starting Point

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

Towards a Language of Description

The theory of organism and the culturalist foundation of biology

The Term Organism

M. Gutmann and E. M. Neumann-Held

The following from Gutmann (1998).

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

5 Biology as an Organismic Discipline

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

M. Gutmann and E. M. Neumann-Held

The theory of organism and the culturalist foundation of biology

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