The Disappearance of Form? Some Methodological Considerations on a Lost Conceptual Dimension in Biology

Abstract

The concept of form belongs—apparently—to an older stage of biological concept formation. Paradoxically, it is even the insistence on the reference to form that shows its very disappearance.1 The more the functionalization2, systematization, and finally algorithmization of modern biology3 advances, in the sense of systems biology, synthetic biology and bioinformatics, the less audible the call for a rehabilitation of the concept of form becomes.This technomorphic tendency, to deal with living entities in terms of artifacts, increasingly brought the concept of transformation into the forefront—articulated, e.g., in the formal sense by differential equations and their combination on all “levels” of the biological organization of living entities.4 It also coincidentally furthered the skepticism concerning the fundamental difference between artifacts and living entities. By explicitly denying or overlooking the conceptual necessity of this difference, the sensitivity for the peculiarity of living entities also dwindled, which in turn even in the more recent history of biology regularly evoked debates on the autonomy of biology and the “irreducibility” of its objects.5Against this background, the concept of form seems to be dispensable, the more so as it is often associated with substantialism or essentialism. By assuming the description-independent existence of some essence, both approaches address organisms or species as belonging to well-defined, unchangeable natural kinds.6 These “-isms” are suspected of being indicators of vitalist obscurantism, as exemplified by the well-known vis vitalis or entelechy and similar metaphysical monsters. In particular, Neo-Aristotelian approaches came under severe criticism7 for being major obstacles to transforming biology into a kind of modern science, because those approaches seemed to be rather susceptible to metaphysical elements.8This suspicion is not totally unfounded, insofar as even in more recent, albeit closed debates of the 1970s and 1990s, it was structuralist, system- and complexity-theoretical concepts9 that demanded form as relevant in a way, which was often unacceptable for mainstream research.10The main focus was on the juxtaposition between form-oriented approaches, which tended to consider form a specific factor beside or above pure material dispositions on the one side, and particularly developmentalist approaches on the other side, which dealt with form as an empirical problem. However, a third option was regularly overlooked, namely the reference to form as a categorical necessity, the analysis and elaboration of which is the aim of this paper. In order to achieve this goal a functional11 and by no means substantialist understanding of the predicative mode of “substantial”12 statements is provided, which allows the introduction of form as an integral part of the methodological foundation of biology.When we ask for the “form”13 of something, we have to face certain indeterminacies of natural languages. In Greek the two expressions εἶδος and μορφή are occasionally used synonymously; still there is a difference, which might be perceived by associating μορφή with the sensible appearance of something in terms of its shape. In a certain contrast to μορφή, the εἶδος has some epistemic14 connotations to “knowing something” in a very broad sense covering “esthetic” as well as conceptual judgment. The relation between “shape” and “knowledge” explicated by form-concepts, is essential for the focus of the present paper, because the argument aims at the connection between form and its determination by specific modes of descriptions that articulate some specific knowledge and knowhow about living entities.Thus, “having or displaying” a shape, or “being shaped” can be considered an object-level expression of an esthetic judgement15 (e.g., about living entities or artifacts), whereas “form” as a meta-level expression articulates these judgements in terms of explicable and explicit knowledge. In order to gain a starting point, we should therefore discern three relevant grammatically and hermeneutically different ways of dealing propositionally with shape, primarily used for the characterization of artifacts:Something appears predicatively as “having a shape,” in contrast to its negation16, whereby the opposition need not be contradictory, since the state of something thus determined allows for gradations.Procedurally something appears as becoming shaped, if it gains a shape, or comes into it, whereby the transition from “x is unshaped” to “x is shaped” is allowed, but also that from “x is A-shaped” to “x is B-shaped.” In this substantial mode of speech, the utterance may refer to certain (non-lingual) practices that not only accompany the transition but articulate it practically. Accordingly, “to shape” does not appear as a common verb, because we explicate it by reference to other activities—such as the kneading of clay, the improvement of sound formation by training our inspiration—and expiration, or the argumentative structuring of a speech. Even in the realm of living entities this reference can be found when we assume, for example, that a chicken gains its adult shape by “developing itself” from an egg (we will come back to this point). Correspondingly, we refer to “shaping” in the most diverse ways of producing or manufacturing something.17“Shape” is finally applied by referring to something as having been shaped, i.e., “shape” is explicitly the result of a process such as under scrutiny.This differentiation leaves unsolved the problem of whether addressing something as “having” a shape is merely a (subjective) judgement in the eye the beholder or a reference to some “objective” state of affairs—both solutions however are connected to “knowledge” about the respective object. A deeper insight can be gained by replacing the verbal phrase “to shape (something)” by “to generate/produce (something) in a certain way,” a move that allows us to discern reflexive and non-reflexive modes of understanding “shape.” The more familiar non-reflexive case seems to say nothing else but that shape (mode 3) is the result of some productive or generative activity, in that the becoming-shaped appears as in mode 2. This allows even transitive series such as “A generates B,” “B generates C” and thus “A generates C,” as for instance the craftsman, who builds a machine tool that produces tools in virtue of which therefore the craftsman produces the tools. The non-reflexive mode of articulating shaping processes shows an important feature of the generation of artifacts, insofar as the product is not a necessary prerequisite for their actual production: we do not need shoes in order to generate more shoes. The shape then belongs to the thing produced insofar as it is imposed upon the educts; thus “being shaped” occurs in the predicative, determinative mode.18The reflexive use in contrast aims directly at the circumstance that the presence of a shaped entity is already required, as “man begets man” (ἄνθρωπος γὰρ ἄνθρωπον γεννᾷ19); something which is essentially shaped (by “itself”20) will produce something shaped. In contrast to artifacts, living entities cannot be discerned from their respective being-shaped, which occurs in an attributive21, modifying mode, because the being-shaped of living entities is an intensive expression.22In general, the non-reflexive understanding allows the transition from unshaped to shaped entities and should predominantly apply to artifacts and their production, whereas the reflexive understanding is essential for living beings. The relation between these two modes of speech will be examined later—it should be noted however, that the non-reflexive mode has a certain epistemic23 primacy.So far, we have determined “shape” in different modes of utterances as an object level expression without relevant reference to biology at all. Even our application on living entities does not, because we understand this term as belonging to an everyday lifeworld language game. When we shift to biology, we enter a scientific language-game, which differs fundamentally in certain respects. According to Heidegger’s statement “So ist denn auch das Wesen der Technik ganz und gar nichts Technisches” (2000, p. 7)24, it can be said that the essence of biology is nothing biological either. Therefore, if we ask how the objects25 of biology come into existence, we usually refer to some specified knowledge and knowhow that is tightly connected to the mode of questions we assume to be answered biologically. The important thing here is that transitions from everyday lifeworld to biology are actually made from a methodical starting point, according to which something is already known in a certain way (yet not biologically) as something.Aristotle provides some relevant clues for this task, by asking for causes (α ἰτι´α), which are tightly connected to specific modes of questions26 that should allow an object to be established as intelligible in the first place.27 This argumentative move is an indispensable prerequisite for every scientific effort, because explananda are not just given. They must be described in a certain way in order to provide something as something that thus appears as the described explanandum, being “in question”.28Going from questions towards answers, different linguistic and non-linguistic means are applied in order to describe and structuralize the things questioned in such a way that the answer can be given satisfactorily. If we assume that science consists, at least to a certain extent, in giving and taking reasons - λόγον διδόναι και´ί ἀποδέχεσθαι - the famous four Aristotelian “causes” would be just such basic modes of questioning. They lead us from—here—biotic entities, given in a context of common knowledge and knowhow towards scientifically—here biologically-structured abstractions. Aristotle gives a good impression of those questions, which render objects, given in ordinary experience, accessible to scientific research. His, however restricted, research program starts with the question for the “what,” as already existing, that enters into research (τòὸ ἐξ οὗ γι´ίγνεται´ί τι ἐνυπάρχοντος). Here we find what was announced as a “substantial” speech mode29, being logically of a predicative nature. The second question asks for the “form” of something, which explicitly includes examples (τò εἶδος καὶ τò παράδειγμα), followed by the “where from” the change or the rest comes into being (ὅθεν ἡ ἀρχη`ὴ τη˜ς μεταβολη˜ς ἡ πρώτη ἢ τη˜ς ἠρεμήσεως) and finally the “goal” of the process in the sense of the “why” (τò οὗ ἕνεκα) (Physica, 194b 24 ff.).Setting aside the question for the completeness of the causes, the result of answering on this questioning provides descriptions of something as a scientific object, a move that makes the actual investigation possible in the first place. Questions of these and similar types enter into the methodological starting-point, which thus articulates the epistemic interest as well as it provides the descriptive means to obtain it. This epistemic interest is explicated by asking questions referring to peculiar capacities or features of living entities given in an ordinary context. E.g., observing a crayfish performing its tail-flic reflex may give rise to the question how the driving forces for this striking mode of motion are produced, how the force generating elements are connected to the exoskeleton, how the mechanical coherence of the entire living entity is preserved under the massive impulse, etc.30 The same technomorphic procedure underpins questions for how the membranes of neurons produce electric currents, how information is processed in the pre-frontal cortex or how the biomechanics of the supporting structures of tetrapod vertebrates can be understood.31By answering questions of this kind (how is x performed, where does y come from, of what kind is z, etc.) we are leaving the epistemically reduced frame of ordinary language and everyday experience. The underlying difference between living entities and biological objects can then be determined as follows:With the term “living entities” we refer to essentially everyday lifeworld practices; in this speech mode we represent biotic entities, that do not have the property of being alive, but whose essence consists in being alive. This implies that “Being” is in this use an intensive expression, too.32In contrast to this enthymematic speech-mode, technical descriptions are applied in order to shift from living entities to their organismic representations. An ὄργανον33 is a tool and accordingly, organisms are functionally described and structured living entities, or more precisely, living entities being described as if they were functional units, which we will refer to in the following considerations as organisms or organismic constructions.34 This process usually starts with (morphological) individuals, that became preparatively35 structuralized by identifying those structures, which are relevant in order to explain the feature or capacity in question as indicated above.As such, organisms, in contrast to the original living entities, consist in structures; they are also formed from them, but not reproduced by them. Against the background of this distinction, life can be understood in two ways, which are determined by the change of language level. While the intensive use of “life” (in the sense of being-alive) provides us with methodological starting points for biological descriptions, the term “life” within the scientific language of biology becomes a concept of reflection (Reflexionsbegriff).36 The original intensive expression refers to enthymematic, empractical knowledge and knowhow; it is a second-order capability enabling living entities to act accordingly, for example as being able to breathe, to grow, to move, to propagate, etc. In contrast, the concept of reflection is tightly connected to scientific theories and articulated by them. It does not have a typical material reference, but refers to a list with different entries, including the usual candidates required by the respective living entity in biological description, like metabolism, respiration, locomotion, development.37 This shift from ordinary to scientific language is accompanied by a conceptual shift in dealing with the respective living entities. The term “form” then denotes an abstract concept that entails the knowledge and knowhow applied to structuralize living as functional units. “Functionality” in this sense is neither connected to nor restricted on “functional or constructional morphology”. Even enzymes, genes, protein domains or transcription factors display functionality in a categorical sense.38 The resulting organismic construction (the organism) can itself be understood as an element of a system, or as a system of elements.39Therefore, not only different questions can be asked40 but also iterated questions of the same type.41 This methodological sequence, both from the organism into a super-organismic as well as into a sub-organismic context, is regularly presented biologically as ontic stratification - for example, from the morphological individual to the organ, tissue, cell, organelle, etc. or vice versa via populations to ecosystems, etc.42So far, the term “form” indicated not only a shift of language level but a shift of practice, procedures and argumentative framing. We introduced the term as an abstract expression, defined in reference to the structualisation of a biotic entity as functional units, named organisms. This is a first—conceptual—step, which mirrors Darwin’s use of breeding as a model for the introduction of central evolutionary expressions.43 Into this procedure of object-constitution enters prescientific, empractical knowledge and knowhow44 as well as other types of (non-biological) knowledge, coming from different sources such as physics, chemistry, technology etc.45This process of object-constitution is however only a first step, which leads into biological conceptualizations of living entities. In fact, we transcend the starting point by not only referring to (morphological) individuals, but to other kinds of processes, which constitute the wider contexts of biotic activities. Thompson (2008) indicates the peculiarity of this wider context as “natural history judgements,” which we want to explicate more exactly by discerning three levels of organismic form as types of reproduction (2008)46:The individual form refers to the organismic structures and their respective bionomic functionality within the framework of the morphological individual. The activities of such individuals determine the mode of being alive in terms of non-extended identical reproduction, insofar as the form provides the invariants for the description of the changes, which the structures of the organism undergo. During ana- and kata-bolism, metabolic regulation etc. organismic individuals remain the same by, through and in spite of changing.The developmental form of individuals can be understood as non-extended non-identical transformation. The invariants of this type of reproduction become the variants within the life cycle of the reproductive units of those forms, which is usually called individual development, as in embryological differentiation, growth, regeneration of limbs etc.The evolutionary form can finally be understood as extended non-identical reproduction, for which the invariants of 1 and 2 become the variants of the transformation.It may become clear now to what extent the shift from a substantial to a functional understanding of being-alive already articulated the concept of form as categorically structuring transformative processes. For we encounter the individual form within the biological theory as an invariant of processes, which allows us to determine its being alive as reproduction of the first type, that is as a (constantly ongoing) maintenance of functional states.With shifting to the developmental form these invariants become variables of a unit, which we (biologically) address as the individual form’s life-cycle articulated in typical propositional statements such as “the chicken develops (itself) from the egg.” According to Aristotle we can rephrase, that the egg—which comes from an adult chicken—is (in the context of this statement) first in time, but ultimate in concept; the chicken that hatches from it, ultimate in time but first in concept.47 More generally, the description of development is a representation of processes that displays a specific categorical structure.The developmental form shows all peculiarities of the reflexive mode; however, it is by no means the chicken which develops “itself” from the egg, nor is it the egg. This well-known riddle becomes solved, by explicating the reflexive mode with reference to the non-reflexive understanding the generation of the developmental result in terms of production. The connected stages of transformation, that constitute non-identical reproduction, are then conceptualized as processes in terms of material and energy transformation.48 Their causal coherence however has to be extended to the entire life cycle, within in which identical reproduction and its constitutive invariants becomes a mere transitory state of the comprehensive unit of the life cycle.The evolutionary form can be determined by a specific speech mode—such as the individual form of organisms and the developmental form of life cycles. As in identical and in non-identical reproduction, the starting point for reconstructing evolutionary transformation is articulated by questions, which aim at the actual lifeworld. It is this lifeworld that becomes the subject matter of questions like “in what way is the peribranchial apparatus of enteropneusta evolved,” or “how does the Coelom evolve in the vertebrate phylum.” As we are here only interested in the form of the answer, which constitutes the logical structure of evolutionary statements, we are starting with what is known best to us (γνωριμωτέρων ἡμι˜ν)—the given life world—aiming at what is to be known best by principle—the ἀρχή or principium, hence the γνωριμωτέρων ἁπλω˜ς or τη˛˜ φύσει.49 In consequence, the extinct lifeworld is reconstructed in the light of the actual lifeworld, the explanation of which however is accomplished by referring to the reconstructed extinct lifeworld. The present situation is thus understood—retroductively50—as the result of transformations of a past, which itself is approached by knowledge acquired in dealing with the present. This results in a peculiar dialectics as the reconstruction reaches from the present into the past, its representation however from the past to the present.This necessary reversal of the order of reconstruction into the order of (re)presentation of the reconstruction, necessitates the reversal of the methodical starting point into the temporal conclusion (the actual lifeworld), and vice versa of the methodical conclusion (the final step of the evolutionary reconstruction) into the temporal starting point. Both movements together constitute the complete form of an evolutionary explanation, according to which the transformation of evolutionary units is reconstructed and presented as a narrative on the origin of the respective forms.51Evolutionary reconstructions thus build a meta-level to functional biology in the broadest sense—including individual as well as developmental forms. However, from a methodological point of view, this meta level is not just a mere shifting of the time scale.52 For not only does the (functionally characterized) past state of the lifeworld become a mere generic precondition of the actual lifeworld, being the result of the latter’s functional analysis and subsequent reconstruction. Rather, the actual lifeworld itself is now conversely considered to be a mere case of the reconstructed extinct lifeworld being its transformative result.The categorical foundation of biology finds its methodological conclusion in evolutionary-theoretical reflection on the principles of transformation of living entities on all three levels of reproduction.53 According to the extent in which form is either understood merely as an empirical problem or as a dispensable remnant of pre-modern natural history that would have to be removed in toto from biology, the sensitivity for the difference between categorical and empirical problems is simultaneously lost. This loss of the categorical basis of modern biology has its origin in the loss of form as well as in the loss of the form of forms, which both belong to the categorical foundation of biology.Our diagnosis of the disappearance of form seems to have been premature—at least when we are willing to answer the question for necessary prerequisites and anticipations even of pure empirical biological research. In order to do so, we reflected on the logical structure of the being-shapedness of living entities, which revealed a necessary connection to different types of knowledge. Introducing organisms as deep-structured conceptual representations of living entities, we exposed three levels of transformation that rule the biological ontology in terms of individual, developmental and evolutionary form.These types of form-thinking eventually allow us to shed some light on the specific epistemic situation of the reconstructive evolutionary biologist. Evolution presents itself as a perfect example of a self-referential historical process. It started a few billion years ago with the first “living” entities (of whatever sort). It led to enormous diversification and differentiation (of whatever structure) and eventually generated our recent lifeworld (not yet known and understood in detail and in its entirety), the reconstruction of which became the subject of the evolutionary biologist, culminating in the epistemological analysis of the reconstructing biologist as such. This self-referential structure is the very form of a hermeneutic reflection, which necessitates the analysis of the inherent temporal logic of human historicity54—a task that presupposes the categorical form of form-thinking, we explicated in this paper.