The unsolved problem of the creation and inheritance of organismic forms In his article: “Radical Constructivism in Biology and Cognitive Science” John Stewart wrote: “In Mendelian genetics1, the "genotype" as such cannot be observed directly2: it is a theoretical hypothesis or postulate3. Differences in genotype are inferred from observed differences in a phenotypic character; if there are no phenotypic differences, no genotypes can be inferred. It follows that there is not and cannot be any genetics of a character which is invariant in the species in question. Thus, genetics can and does explain the distribution of (for example) blue eyes vs brown eyes in the offspring of particular crosses; but if all the animals in the population have eyes, genetics does not and cannot explain why there are eyes in the first place that might be blue or brown. To take another example, genetics does not and cannot explain why the offspring of cats are kittens which grow up into cats that resemble their parents. This is the sense in which genetics is not a science of heredity. The "blind spot" is even more complete if the character in question is common to all living organisms. Thus, the phenomenon of autopoiesis - the fact that all living organisms produce themselves by their ongoing dynamic processes, which we shall examine more closely below - is and can only be ignored by Mendelian genetics and, hence, by the neo-Darwinian paradigm. It is to be noted that this is a true blind-spot, in the sense that not only is there a key feature of living organisms that is not seen; from a neo-Darwinian perspective, it is intrinisically impossible to appreciate that there is a feature which is not seen. Nevertheless, this lacuna gives rise to a serious weakness in the neo-Darwinian account of evolution.” 1 In what follows, I shall use the simple term "genetics" instead of referring pedantically each time to "Mendelian genetics". Note, however, that this carries a risk of confusion: Mendelian genetics is precisely not a science of the constructive processes of genesis, as in "phylogenesis", "ontogenesis" or Piaget's "genetic structuralism". 2 Nowadays, of course, DNA can be observed; but the nucleotide sequence of a piece of DNA is not a genotype, and to confuse the two is a reification which generates grave conceptual confusion. 3 This is not an objection: in physics, all the fundamental scientific objects, such as atoms, quarks, and electro-magnetic fields, cannot be directly observed as such. It should, however, as a brake on the slippery slope to objectivist reification. In this context I also may quote Denis Noble “Neo-Darwinism, the Modern Synthesis and selfish genes: are they of use in physiology?” J. Physiol. 589.5 (2011) 1007–1015: “As I have argued elsewhere (Noble, 2008) the main difference between DNA and nonDNA inheritance is simply that one is digital, the other analog. In developing the organism the 3D analog information is just as necessary as the 1D digital (DNA) information. Neither is sufficient by itself… Moreover, organisms are not in fact digital machines (Shapiro, 2005; Noble, 2010a).” In these two articles the authors refer to a fundamental problem of Biology. Namely, there is no molecular-biological explanation of how the form of an organism is maintained (or altered in a potentially useful way) in the constant metabolic flux of structural modifications. If one does not know that, one cannot explain how a certain form is inherited. The suggested dependence of a phenotype on a genotype has no sound scientific foundation, if we infer a molecular basis for the genes in the sequence of nucleotides (apart from the problem that it is difficult to characterize the nature of the genotype; I may quote Noble in the article above: “The concept of a gene has changed, and is still changing, so what version do we use?”). The alleged relation of the phenotype on the genotype is also misleading because it is based in an unjustified generalization of Mendelian genetics, in which differences in a phenotype are identified with the phenotype as a whole (see Stewart, above). For this reason we have to answer the question how the analog information of a form, mentioned by Noble, is stored and inherited. Genetics does not give a satisfactory answer, as has been stated by the geneticist Stewart. Inheritance of specific forms of a species is therefore still an unsolved problem in Biology. I tried to solve this problem by transferring Whiteheads idea of the interdependence of actual entities (in which the outcome of antecedent actual entities is “inherited” by subsequent actual events) to a chronology of intracellular adaptive events. A molecular-biological explanation of inheritance of a certain organismic form is impaired by the fact that it is impossible to obtain a molecular model of morphogenesis, for the following reason: Firstly, the form of an organism A is what is perceived by an organism B. If, for example, we could somehow experience the electric field of a fish, we would perceive this organism in a different manner. A molecular model of how we see the form of an organism must therefore be loaded with the properties of our highly developed perceptive apparatus. Since the form is not an objectively given entity which is independent from an observer, morphogenesis cannot be derived from the metabolic activities. Naturally this problem is a consequence of the difference between organisms and non-living beings. In the latter case the form is the result of the arrangement of components, whereas in organisms the structure is constantly altered although the form is maintained. This means that in organisms the form determines how the components of its different structures are permanently re-arranged during its development and not vice versa. In this process a cell or an organism strives for functional harmony of all its partial processes. Morphogenesis has therefore a ‘geistige’ (cannot be translated, perhaps psychophysical?) dimension in the sense that it is related to an ideal. This dimension does not exist in chemistry. Secondly, there is the problem of ‘autopoiesis’ addressed by Stewart in the quotation above. To explain this, let us consider the proliferation of the spindle-shaped green alga Ankistrodesmus. In this case a mother cell divides into several spindle-shaped daughter cells, the socalled ‘autospores’. Then the cell wall is degraded and the little spindle cells are released. In the subsequent growth process in which each of the autospores finally becomes a new mother cell, the shape of a spindle is maintained. The form of this alga is determined by the cell wall, which is located outside of the cell membrane. For this reason during growth of the cell the biosynthesis on each position of the cell wall must be coordinated with the biosynthesis on all other positions, because otherwise the form of the cell would be distorted. Furthermore, the biosynthesis of the cell membrane, the cytoskeleton etc. must be also conformed to the enlargement of the cell wall. During growth all biochemical partial process must therefore constantly and simultaneously be adjusted to the recreation of that particular form, since an increase of the cell volume requires a permanent reconstruction of its structures. But this means that the constantly occurring establishment of a certain form during the development of a cell has a teleological aspect. Thereby, a not yet existing final state regulates the coordination of all partial processes for re-attaining the intended form by the organism. Such a coordination, endowed with an anticipatory orientation towards the future, requires the existence of an internal relationship among all partial processes. For this reason a coordinating principle is a precondition for maintaining the certain form, by which the cell distinguishes itself as a unity from its environment. Chemistry does not provide such a coordinating principle and therefore is insufficient for an explanation of morphogenesis. In order to deal with this problem I have previously suggested (in our discussion with Salthe) to consider the organismic nature as a correlate to technical practice that comprises a producer and its product. The products are the structures, by which the organism constantly re-creates a potentially useful form during its development. The structures can be analysed and localized in objective terms. But the producer is a precondition for the creation of objective structures and therefore, as the creative organismic subject, evades an objective description. Since there is an ontological difference between a producer and its products, the producer can be found in the cell by searching for an ontological difference in metabolic activities. In a previous publication (in “Beyond Mechanism, Putting Live Back into Biology”) I have explained how this can be achieved by analysing interplay of energy converting processes in the cell. Setting out from this physiological explanation I have then (inspired by John Dewey) postulated that the energy flow through a living system serves as a medium for a ‘field of tension’, involved in anticipatory information processing about a potentially useful reconstruction of energy converting subsystems. In this process, organismic identity is based on an internal representation, provided by this ‘field of tension’ that emerges self-referentially from and acts upon reconstruction of energy converting subsystems. Thus, such a ‘field of tension’ carries information about the form of a species in the individual development and in a succession of generations. Therefore the ontological difference between a ‘field of tension’ representing a general feature of the organism and its partial processes guides the process of morphogenesis. The ontological difference is revealed in interplay between nascent form and attained physical form (again better in German: between werdender Form und gewordener Form; for Ernst Cassirer between forma formans and forma formata). In its form an organism gives itself a physical expression by which it tries to exert pressure on its surroundings. In higher developed organisms this is also the basis for bio-communication: in this case an interpretation of the expression of an organism A by an organism B determines how B expresses itself in response to this and vice versa. From the sequence of mutual interpretations of acoustic expressions finally language emerges and, as a result of the collective reflexivity of the language, higher forms of consciousness (for details of this process see: Ernst Cassirer, “Philosophy of Symbolic Forms). Along this line I tried to develop a ‘theory of organisms’, in cooperation with the Swiss process philosopher Reto Luzius Fetz. It will be published in near future. It is based on some thoughts of Whitehead, but in order to accomplish this we had to adjust the categoreal scheme of Whitehead to Biology.