Quaderni del Centro Interdipartimentale L. Galvani 2

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1 Quaderni del Centro Interdipartimentale L. Galvani 2

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3 IMMUNOLOGY TODAY Three Historical Perspectives under Three Theoretical Horizons Edited by Andrea Grignolio

4 This book has been published with the support of the Fondazione Sigma-Tau. Questo volume è stato pubblicato con il contributo della Fondazione Sigma-Tau. centro interdipartimentale l. galvani alma mater studiorum università di bologna Bononia University Press Via Farini 37, Bologna tel. (+39) fax (+39) Bononia University Press Tutti i diritti riservati ISBN Progetto grafico di copertina e impaginazione: DoppioClickArt Stampa: Editografica Prima edizione: marzo 2010

5 To Francesca, who brought love and theatre into my life.

6 Frank M. Burnet believed passionately that, while advanced techniques were essential tools, science was primarily about ideas. His extraordinary gift was to take apparently unconnected observations and fit them, almost force them, into whatever theoretical approach was his current obsession. Sir Gus Nossal, The Lancet, 1985 I find it astonishing that the immune system embodies a degree of complexity which suggests some more or less superficial though striking analogies with the human language. Niels K. Jerne Nobel Lecture, 1985 The immune system is, like the brain, an organ of adaptation to external environment, recording biochemical instead of sensory experiences. Like the brain, the immune system is said to have a memory David W. Talmage The Nature of the Immunological Response, 1969 practically all molecules in the universe are antigens. Edward S. Golub Immunology: A Synthesis, 1987

7 Contents Introduction 9 Claudio Franceschi, Andrea Grignolio PART I: HISTORY OF IMMUNOLOGY A Tale of Two Immunologies 15 Alfred I. Tauber Immunology: A Historical Perspective 35 Gilberto Corbellini Networks, Degeneracy and Bow ties: A Unifying Perspective of Immunological Paradigms and Architectures 53 Andrea Grignolio, Paolo Tieri, Gastone Castellani PART II: HISTORY OF THE IMMUNE SYSTEM The Common Origin of Immune and Neuroendocrine Systems 71 Enzo Ottaviani, Davide Malagoli Immunosenescence within an Evolutionary Perspective 79 Claudio Franceschi, Andrea Grignolio PART III: IMMUNOLOGICAL BIOGRAPHY Flow Cytometry and the Complexity of the Immune System 103 Andrea Cossarizza Role of B cells in the Induction of Peripheral T-cell Tolerance 111 Francesca Granucci, Ivan Zanoni

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9 Introduction This book collects the proceedings of the scientific event Immunology Day which took place in Bologna on May 7 th 2008, and is published as the second volume of the series Quaderni del CIG sponsored by the Sigma-Tau Foundation, the Interdepartmental Center Luigi Galvani Interdepartmental Center for Bioinformatics Biophysics and Biocomplexity (CIG) at the University of Bologna, the European Project Acume2 Interfacing Science and Humanities and the Institute of Advanced Studies (ISA) at the University of Bologna, Italy. On that occasion, the latter assigned to Alfred I. Tauber, a philosopher of science and an immunologist, the ISA Medal for Science, the highest honour awarded by the Institute in recognition of research contributions characterized by both excellence and influence. The small group of scientists who took part in this event was asked to address some hot topics in immunology by focusing their contributions on the idea that further progress would be possible only through the attempt to go beyond the narrow boundaries of an academic vision and to accept the challenge of contaminating immunology with other disciplines in a more integrated and comprehensive approach.

10 10 Introduction Already in the first volume dedicated on Biocomplexity at the Cutting Age of Physics, Systems Biology and Humanities we tried to cross the borders between the so-called two cultures, suggesting that biocomplexity could be assumed as a useful paradigm and new hermeneutical tool also for the humanities. With this second volume we are still on this track, continuing our methodological discourse on the fecundity of travelling from one discipline to another. The fil rouge of this book is that immunology is a highly developed and sophisticated discipline, with a long tradition of theoretical debates, that once again can be the forerunner to test some of the most urgent questions in biology and medicine. Accordingly, most of the contributors share the premise that a historical perspective is essential not only to interpret the available results but also for the immunology of the next future. As a matter of fact, in their essays they articulated the historical dimension on three different levels: 1. THE history of Immunology as a (central) tool that can help improve our epistemological awareness of disciplinary problems (see part I); 2. THE history of the Immune System as an evolutionary point of view essential to understand its present status and developmental processes (see part II); 3. THE Immunological biography, i.e. immunological history at the individual level (ontogenetic time), which is probably the best scientific approach to study the epidemiology and the pathological onset and progression of complex immunological diseases such as autoimmune diseases as well as of age-related pathologies (see part III) which have a strong immune-pathological component (atherosclerosis, diabetes, dementia, among others). Most importantly, these three historical dimensions explain and are explained by three different theoretical horizons, i.e. Degenerated/Bow-tied Architecture, Systems Biology, Ecoimmunology. A few explanatory words may serve to elucidate how the two levels of analysis could work together to provide a unified view of the emerging trends of today s immunology. In biological systems, Degeneracy is generally defined as the ability of elements that are structurally different to perform the same function (many-to-one or one-to-many logic). Degeneracy is a pervasive characteristic of living organisms and it is present at all levels of biological organization. The heuristic value of addressing the immune system as a complex system and under the perspective of degeneracy is that we are forced to imagine operative/architectural structures that can do the (difficult) job of conveying a variety of information on the same entity (a receptor or a cell), being able to integrate such a complex variety of signals in order to have a fine tuned output/response(s) which, in turn, would be further treated in the same (integrated) way by another entity in the system. According to recent suggestions, the so called bow-tie structure is discussed in the book as a reason-

11 Introduction 11 able candidate for the operative/architectural structure which could represent the new unit of the immune system as well as of other systems in the body. We are aware that these arguments will definitely be improved and changed but we have the feeling that we are on the right track even without definitive answers. It is quite fashionable nowadays to talk about Systems Biology relatively to everything, and the fact that science follows fashionable topics is well known. However, looking more closely, the real Systems Biology is far from being a reality and it is more an intuition than a well documented fact. In any case, we agree that a systemic approach to the major problems of Biology and Medicine, such as aging and complex diseases among others, is becoming more effective and increasingly popular. Thus, it is fair to say that Systems Biology is in its infancy and we have to help it grow and became an adult science. This perspective is envisaged in the present book where a systemic approach to immunology is predicted and where some possible examples are illustrated. In other words, the Systems Biology approach is urgently needed to put the vast available knowledge of the immune system within a unitarian framework and to understand its challenging complexity. Ecology is one of the most interesting but complicated sciences and it is a pity that most immunologists are not familiar with such a discipline, which has a lot to teach to immunology. Indeed every animal which has to cope with an invasion from an infectious intruder and has to trigger an immune response will spend a lot of energy on such an action. However, the rule of energy saving is also true for immune responses, and it is clear that a strong immune response will subtract energy for other critical purposes such as movement and reproduction. These considerations are at the basis of a new vision of immunology which takes into account not only the minimization of the cost of immune responses but also the fact that they occur in specific ecological niches, a fact that will eventually decide about the efficacy of the immune response itself. ECOIMMUNOLOGY, too, is in its infancy and there are very few studies in humans and other mammals. A main reason is that humans are very difficult to study even from this point of view and that other animal models such as rodents are usually kept in artificial environments, which are hugely different from the natural conditions which are present in the wild. Despite these limitations, an ecoimmunological approach is urgently needed if we wish to study the immune responses in a broader and more realistic perspective. In the book some contributors argue that some of the evolutionary negotiations which established the rule of the immunological game towards infectious agents within specific environment can became detrimental in the last part of life (aging and senescence), a post-reproductive period largely uncommon in the wild in which the force of selection fades away. It is interesting and important to say that for the first time we provide tools that can help us in applying such a comprehensive theoretical and experimental approach to immunology. We are starting to apply high-throughput technologies

12 12 Introduction with the huge power of all the omics (genomics, transcriptomics, epigenomics among others) to the immune system. Together with the huge computational power available in many labs and with the help of bioinformaticians and mathematicians we are confident that a new era is coming and that immunology (like many other sciences) will be re-visited and re-written according to these new systemic approaches. Claudio Franceschi and Andrea Grignolio Bologna October 2009

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