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This is the invited evening lecture of the biannual workshop on hydroid development of 1999. Its topic is the role of hydra as a rather puristic model for the de-novo generation of spatial patterns in development, and our work in this field. Emphasis is placed not only on experimental studies, but also on theoretical analysis, because the understanding of spatial order requires a systems approach involving the combination of knowledge on molecules, cells and tissues with mathematical analysis, laws and facts.
Understanding cooperative human behaviour depends on insights into the biological basis of human altruism, as well as into socio-cultural development. In terms of evolutionary theory, kinship and reciprocity are well established as underlying cooperativeness. Reasons will be given suggesting an additional source, the capability of a cognition-based empathy that may have evolved as a by-product of strategic thought. An assessment of the range, the intrinsic limitations, and the conditions for activation of human cooperativeness would profit from a systems approach combining biological and socio-cultural aspects. However, this is not yet the prevailing attitude among contemporary social and biological scientists who often hold prejudiced views of each other's notions. It is therefore worth noticing that the desirable integration of aspects has already been attempted, in remarkable and encouraging ways, in the history of thought on human nature. I will exemplify this with the ideas of the fourteenth century Arab-Muslim historian Ibn Khaldun. He set out to explicate human cooperativeness - "asabiyah" - as having a biological basis in common descent, but being extendable far beyond within social systems, though in a relatively unstable and attenuated fashion. He combined psychological and material factors in a dynamical theory of the rise and decline of political rulership, and related general social phenomena to basic features of human behaviour influenced by kinship, expectation of reciprocity, and empathic emotions.
Full applicability of physics to human biology does not necessarily imply that one can uncover a comprehensive, algorithmic correlation between physical brain states and corresponding mental states. The argument takes into account that information processing is finite in principle in a finite world. Presumbly the brain-mind-relation cannot be resolved in all essential aspects, particularly when high degrees of abstraction or self-analytical processes are involved. Our conjecture plausibly unifies the universal validity of physics and a logical limitation of human thought, and it does not regard consciousness -the most basic human experience - as a marginal phenomenon. ++++ RATIO appeared up to 1987 in both a German and an English version. The German title of this article: Alfred Gierer, Der physikalische Grundlegungsversuch in der Biologie und das psychophysische Problem. RATIO XII, Heft 1, 1970, S. 40-54.
Modern brain research related to consciousness has resulted in many interesting in- sights, for example into the neurobiological basis of attention and of language. In biological terms, human consciousness appears as a system’s feature of our brain, with neural processes strictly following the laws of physics. This does not necessarily imply, however, that there can be a general and comprehensive scientific theory of consciousness. Predictions of the extent to which such a theory may become possi- ble vary widely in the scientific community. There are reasons - not only practical but also epistemological - why the brain-mind relation may not be fully decodable by finite procedures. In particular, analogies with mathematical theorems of un- decidability suggest that self-referential features of consciousness, such as multiple self-representations like those involved in strategic thought, may not be fully resolv- able by brain analysis. Assuming such limitations exist, this implies that ob jective analysis cannot exhaust sub jective experience in principle. A person’s consciousness and will are accessible to external observation only within limits. In some respects, we do not even learn to know ourselves except by our actions. It thus appears that a scientific look at consciousness and the human mind, combining universal physi- calism with epistemological scepticism, is not inconsistent with certain concepts of sub jectivity that are current in the humanities, despite all the differences in the style and terminology of discourse.
Socioeconomic inequalities are functions not only of intrinsic differences between persons or groups, but also of the dynamics of their interactions. Inequalities can arise and become stabilized if there are advantages (such as generalized wealth including “human capital”) which are self-enhancing, whereas depletion of limiting resources is widely distributed. A recent theory of biological pattern formation has been generalized, adapted and applied to deal with this process. Applications include models for the non-Gaussian distribution of personal income and wealth, for overall economic growth in relation to inequalities and for effects of uncoupling strategies between developing and developed countries. Note added after publication: The equations (14) for the model of the income distribution, with its characteristic non-Gaussian extension towards higher incomes (fig.4), are closely related to the Fokker-Planck equation that is widely applied in many fields of physics.
The introductory personal remarks refer to my motivations for choosing research projects, and for moving from physics to molecular biology and then to development, with Hydra as a model system. Historically, Trembley’s discovery of Hydra regeneration in 1744 was the begin¬ning of developmental biology as we understand it, with passionate debates about preformation versus de novo generation, mechanisms versus organisms. In fact, seemingly conflicting bottom-up and top-down concepts are both required in combination to understand development. In modern terms, this means analysing the molecules involved, as well as searching for physical principles underlying development within systems of molecules, cells and tissues. During the last decade, molecular biology has provided surprising and impressive evidence that the same types of mol¬ecules and molecular systems are involved in pattern formation in a wide range of organisms, including coelenterates like Hydra, and thus appear to have been “invented” early in evolution. Likewise, the features of certain systems, especially those of developmental regulation, are found in many different organisms. This includes the generation of spatial structures by the interplay of self-enhancing activation and “lateral” inhibitory effects of wider range, which is a main topic of my essay. Hydra regeneration is a particularly clear model for the formation of defined patterns within initially near-uniform tissues. In conclusion, this essay emphasizes the analysis of development in terms of physical laws, including the application of mathematics, and insists that Hydra was, and will continue to be, a rewarding model for understanding general features of embryogenesis and regeneration.
The short paper introduces the concept of possible branches of double-stranded DNA (later sometimes called palindromes): Certain sequences of nucleotides may be followed, after a short unpaired stretch, by a complementary sequence in reversed order, such that each DNA strand can fold back on itself, and the DNA assumes a cruciform or tree-like structure. This is postulated to interact with regulatory proteins.
Aggregates of previously isolated cells of Hydra are capable, under suitable solvant conditions, of regeneration forming complete animals. In a first stage, ecto- and endodermal cells sort out, producing the bilayered hollow structure characteristic of Hydra tissue; thereafter, heads are formed (even if the original cell preparation contained no head cells), eventually leading to the separation of normal animals with head, body column and foot. Hydra appears to be the highest type of organism that allows for regeneration of the entire structure from random cell aggregates. The system is particularly useful for studying cell interactions, tissue polarity, pattern formation, and cell differentiation.
The paper addresses the formation of striking patterns within originally near-homogenous tissue, the process prototypical for embryology, and represented in particularly puristic form by cut sections of hydra regenerating a complete animal with head and foot. Essential requirements are autocatalytic, self-enhancing activation, combined with inhibitory or depletion effects of wider range - “lateral inhibition”. Not only de-novo-pattern formation, but also well known, striking features of developmental regulation such as induction, inhibition, and proportion regulation can be explained on this basis. The theory provides a mathematical recipe for the construction of molecular models with criteria for the necessary non-linear interactions. It has since been widely applied to different developmental processes.
The generation of viral mutants in vitro was demonstrated by treatment of the isolated RNA of Tobacco Mosaic Virus by nitrous acid. This agent causes deaminations converting cytosine into uracil, and adenine into hypoxanthine. Our assay for mutagenesis was the production of local lesions on a tobacco variety on which the untreated strain produces systemic infections only. A variety of different mutants are generated in this way. Quantitative analysis of the kinetics of mutagenesis leads to the conclusion that alteration of a single out of the 6000 nucleotides of the viral RNA is sufficient for causing a mutation.