sign in sign up
<<
Home , Aernout Mik, Britton Chance, Carel Visser, Charles Weissmann, Daan van Golden, David de Wied

Royal Netherlands Academy of Arts and Sciences

Heineken Lectures 2004

Dr A.H. Heineken Prize for Environmental Sciences Simon A.Levin Heineken Lectures 2004

Dr A.H. Heineken Prize for Environmental Sciences Professor Simon A.Levin delivered his Heineken Lecture on September 28,2004 at the Royal Tropical Institute in Amsterdam. Royal Netherlands Academy of Arts and Sciences

Heineken Lectures 2004 Amsterdam,2005

Dr A.H. Heineken Prize for Environmental Sciences Simon A.Levin The Heineken Lectures series for 2004 also includes Dr H.P.Heineken Prize for and 2004,Andrew Z. Fire Dr A.H. Heineken Prize for 2004, Elizabeth H. Blackburn Dr A.H. Heineken Prize for History 2004,

To order publications from the Heineken Lectures series for 2004, please contact the secretariat of the Academy’s Communications Department at +31 (0)20 551 0726. Contents

page 6 Preface Willem J.M.Levelt

9 Simon A. Levin and his research

12 Presentation address for the Dr A.H. Heineken Prize for Environmental Sciences Johan Bouma

14 The Ecology of Complexity and the Complexity of Ecology Dr A.H. Heineken Prize for Environmental Sciences Simon A.Levin

26 Heineken Lecture Program 2004

27 Audience and publicity for the Heineken Lectures in 2004

28 Secondary School Project about the Heineken laureates

29 General information

30 List of Heineken laureates Preface

Address by Willem Levelt,President of the Royal Netherlands Academy of Arts and Sciences,on the occasion of the presentation of the 2004 on October 1,2004.

Both in the sciences and the arts, history provides us with a looking glass that helps us to focus on the real pioneers. It is much easier for us now to recognize the epoch-making con- tributions of such pioneers as Huygens and Newton, Lavoisier and Pasteur, the Humboldts or Darwin than it was for their contemporaries.A contemporary of Huygens would have had a hard time telling his lasting wave theory from his failing mechanistic theory of gravity.The buzzing genius of Newton spent much more time on alchemy than on the laws of gravity and optics.Who, then, could distinguish the really lasting contributions of these men from their many dead ends? I am deliberately ignoring the abysmal public condemnations of church or state officials concerning some of the loftiest scientific or scholarly insights of their subjects. Neither will I elaborate on the ideologically motivated, unremitting support by church or state of demonstrably false theories such as Lysenko’s Lamarckianism or present-day church and often state-supported creationism. Systematic disinformation of the general public on the achievements of the arts and sciences is always looming. History’s looking glass cannot be dispensed with. Admittedly, however, history can be slow in its filtering exercise. It took no less than 34 years for Father Mendel’s trailblazing genetic discoveries to become recognized by the scientific community, in fact only after others, in particular Hugo de Vries, rediscovered the same laws.These laws never came to the attention of Charles Darwin, a missed opportu- nity to integrate into evolutionary biology. Here it was the scientific community itself that was to blame. Mendel did publish the details of his experiments and theoretical analysis in the 1866 proceedings of his local scientific academy in Brünn, but nobody took any notice of them. The story is hardly different for the arts. In 1723, the town officials of Leipzig, due to appoint a new Thomas cantor, clearly preferred Telemann and Graupner over Johann Sebastian Bach.We would have known better, wouldn’t we? Similarly, the town council here in Amsterdam took down The Oath-swearing of Claudius Civilis that Rembrandt had painted for the new town hall, rolled it up, and returned the masterpiece to him.This in fact led to its disfigurement, because Rembrandt then had to cut the painting down in order to find another buyer.The council clearly preferred the far less controversial town hall contributions by Flinck, Lievens and Jordaens. Clearly, major achievements in science and the arts are by no means recognized as a matter of course, either within the scientific and artistic communities themselves or by society at large. One major function of awards such as the Heineken Prizes is to breed consensus. But consensus on what? Here prizes can serve quite different purposes.There are awards, such as dissertation prizes, whose function it is to make the scientific community aware of talented upstarts. Clearly, our laureates today are not in need of such career prizes.

6 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 All of them are established experts of great repute in their own professional communities. As a rule, major awards such as the Heineken Prizes are never career prizes.They rather fulfil one or both of two other functions. The first one is to highlight a particular landmark empirical contribution. If the Dr H.P.Heineken Prize for Biochemistry and Biophysics had been around in Georg Mendel’s time, he would no doubt have received it for his 1866 paper, probably in the presence of His Majesty King William III. And the reason would not have been the excellence or even the outstanding of this particular work, but rather the fact that it is fundamental to the field.That is the case for Professor ’s discovery of RNA interference, for which he today receives the Dr H.P. Heineken Prize for Biochemistry and Biophysics. It is also the case for Professor ’s identification of the structure of telomeres and her discovery of the telomerase, which will today be honored by the award of the Dr A.H. Heineken Prize for Medicine. In this respect, these two Heineken Prizes are like the Nobel Prizes for Science, which recognize unique breakthrough contributions. In fact, we are proud to say that in many cases, the juries of our Academy’s Heineken Prizes have been well ahead of the Swedish Academy’s committees in identifying such landmark contri- butions. The second function is to highlight a landmark theoretical contribution. Some scientific contributions are fundamental without being discoveries in the strictly empirical sense. Newton experimentally discovered the spectral dispersion of light. In contrast, his breakthrough theory of universal gravity was not an empirical finding, but a theoretical reformulation of fundamental mechanical physics. Today’s Dr A.H. Heineken Prize for Environmental Sciences recognizes Professor Simon Levin’s contributions to fundamental theory,the theory of ecosystem dynamics. For obvious reasons, however, these two types of landmark contributions, the empirical and the theoretical, rarely appear as pure cases. The experimentalist is always theoretically motivated and the only way for the theorist to stay honest is to remain in close contact with empirical work. The two are inseparably interwoven in the study of history. Professor Le Goff’s theoretical reformulation of medieval history emerges from a host of groundbreaking empirical studies. The Dr A.H. Heineken Prize for History recognizes this innovative two-pronged approach. Works of art are too, in their way,empirical contributions.The artist is a discover- er and each work is, to some extent, an experiment in triggering some intended perspective in the eye of the beholder. Mr receives the Dr A.H. Heineken Prize for Art for his ability to create a contextual perspective on the work of art. Where should such consensus be established? First of all in the professional com- munities themselves. A Heineken Prize tells the laureate’s peers:‘this work is fundamental’. A modern scientific peer community is usually quite able to recognize excellence. But it can still take years before it reaches consensus on which new insights are essential to the blue- print of their science. Second, but equally important, is to reach consensus in the larger community, which cares, or should care, about the contributions of science and scholarship to society.

7 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 As Simon Levin expressed it in a recent interview, ‘Public interest is on the macro scale’. He was, of course, referring to macro scale effects in the environment, such as the mainte- nance of biological diversity,but there is a more general issue here.The public at large is not so much interested in telomerase or RNA interference, but rather in questions such as ‘Will it give us a cure for cancer or for AIDS?’. And here there is a major gap to bridge. Professor Blackburn, in a recent interview, gave the example of Gleevec, an effective treatment of leukemia. There was a 30-year gap between the discovery of the chromosomatic disorder in this type of leukemia and the development of an effective drug.There is, as a rule, no linear pathway from knowledge to treatment. But the gap is even wider than this example suggests. In many cases it is simply counterproductive to go for a cure or an application that is understandably wanted by the general public, for the simple reason that at the outset the scientist doesn’t know what potential knowledge is relevant to the case at hand. Eventually, there is only one way for the scientist to proceed. It is to sit down and dissect the system, whether it is a chromosome, a cell, a layered system in the environment, or a state of affairs in medieval history.The process of discovery is entirely self-governed. It has its own logic.To be successful, it should not be deterred by public pressure, by a push for quick . As Professor Blackburn put it recently,‘We weren’t looking to cure cancer and yet it turns out that the enzyme telomerase is one of the most frequently found characteristics of cancer cells.That was not expected’. At the same time, the scientist has a responsibility to explain this state of affairs to the general public, time and again. Why is it that we are spending public funds in this indirect, detached fashion? The Heineken Prizes invite the general public to regard these laureates as model cases. Each, in his or her own way, has made a major effort to inform the general public, to explain the relevance of their work for our living environment, for our health care and for our understanding of ourselves as human beings. If their outstanding example helps to shape public opinion, these prizes will have been money well spent.

Willem J.M. Levelt President of the Royal Netherlands Academy of Arts and Sciences

8 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Simon A. Levin and his research

The research The Royal Netherlands Academy of Arts and Sciences has awarded the Dr A.H. Heineken Prize for Environmental Sciences 2004 to Professor Simon A. Levin for ‘his insights into the effects of scale on ecosystems’. At the basis of Simon Levin’s work is his use of mathematical tech- niques and models to understand the properties of ecosystems, i.e. the biological communi- ties that inhabit specific areas, including all associated living and non-living factors. Levin unites theory and empiricism, with ‘scale’ as his leitmotiv. Natural populations and their internal coherence are influenced in part by time, space and complexity.Something that occurs on one order of scale may have a very different impact on another. Levin seeks concrete answers to such questions as: when does an eco- system collapse? How resilient is it? What is the value of a single species, and what can the ecosystem do without? What roles do evolution and the play? In other words: what are the dynamics of ecosystems? The answers to these questions are highly important not only for research into but also for environmental protection. Levin has shown that many of the properties of ecosystems vary according to fixed patterns and that we often only understand phenomena when we know what is going on at different orders of scale. His insights have led to fundamental changes in the discipline of ecology,which until the early nineties had been divided into a number of sub disciplines, each of them myopic in its own way.What Levin did was offer a pair of ‘glasses’: his article ‘The Problem of Pattern and Scale in Ecology’, which appeared in Ecology. It became the most highly cited work in the entire field in the 1990s.

The laureate Born in 1941 in the , Simon Asher Levin has conducted research in mathe- matical biology for more than forty years. He began by studying mathematics and took his Ph.D. in this subject in 1964 from the University of Maryland, College Park. In 1965 he became a researcher at Cornell University (Ithaca, N.Y.),where he quickly joined the Ecology and Mathematics department. Levin was appointed Professor of Applied Mathematics and Ecology in 1977, a post he held until 1992, when he left Cornell for Princeton University (New Jersey). Today he is the George M. Moffett Professor of Biology at Princeton and the director of the Center for Biocomplexity. Simon Levin is an exceptionally active scientist.A list of his management positions, editorial posts and lectures, the meetings and conferences he has organised or co-organised, and the Ph.D. candidates and post-docs he has supervised runs to dozens of pages. Added to this is his long list of publications (including the 4,800-page Encyclopedia of Biodiversity,

9 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 of which he is the editor-in-chief). He has been the recipient of many prizes and marks of recognition. For example, in 2000 he was made a member of the American National Academy of Sciences and in 2001 was presented with the Akira Okubo Lifetime Achievement Award. Levin is said to be a gifted speaker and an excellent teacher, capable of inspiring and motivating researchers and making his insights comprehensible for the public. He also influences the international research agenda, for example as the chair of the Executive Committee of the International Institute for Applied Systems Analysis (IIASA), an inter- disciplinary research institute based in Austria which studies the human dimensions of global change.

Key publications Levin, S.A. 1992.The Problem of Pattern and Scale Durrett, R. and S.A. Levin. 1994. Stochastic spatial models: in Ecology, Ecology, 73 (6). a user’s guide to ecological applications. Philosophical Levin, S.A. 1998. Ecosystems and the biosphere Transactions of the Royal Society of London,Series B 343: 329-50. as complex adaptive systems. Ecosystems. 1: 431-436. Durrett, R. and S.A. Levin. 1994.The importance of Levin, S.A. 1999. Fragile Dominion, complexity and being discrete (and spatial).Theoretical Population Biology 46: the commons, Perseus Books Group, Reading, MA. 363-94. Levin, S.A. 2003. Complex adaptive systems: Exploring Flierl, G., D. Grünbaum, S.A. Levin and D. Olson. 1999. the known, the unknown and the unknowable. Bulletin of From individuals to aggregations: the interplay between the American Mathematical Society 40: 3-19. behavior and physics. J.Theoretical Biology 196: 397-454. Levin, S.A. and C. Castillo-Chavez. 1990.Topics in Kareiva, P.and S.A. Levin (eds.) 2003.The Importance evolutionary ecology.pp. 327-58. In: S. Lessard (ed.), of Species: Perspectives on Expendability and Triage. Mathematical and Statistical Developments of Evolutionary Theory. Princeton University Press. 427 pp. NATO ASI Ser. C, vol. 299, Kluwer Academic Publishers, Levin, S.A. 1970. Community equilibria and stability, Dordrecht,The Netherlands. and an extension of the competitive exclusion principle. Levin, S.A. and H. Muller-Landau. 2000.The emergence American Naturalist 104(939): 413-23. Reprinted in: of biodiversity in plant communities. Comptes rendus de Niche:Theory and Application, 1975. l’Académie des sciences ‘Sciences de la vie / Sciences 323: Levin, S.A. 1974. Dispersion and population 129-39. interactions. American Naturalist 108(960): 207-28. Levin, S.A. and L.A. Segel. 1985. Pattern generation in Levin, S.A. 1976. Population dynamic models in space and aspect. SIAM Review 27(1): 45-67. heterogeneous environments. Annual Review of Ecology Levin, S.A. and R.T. Paine. 1974. Disturbance, patch and Systematics 7: 287-311. formation, and community structure. Proceedings of the Levin, S.A. 1978. On the evolution of ecological National Academy of Sciences,USA 71(7): 2744-47. parameters. pp. 3-26. In: P.F. Brussard (ed.), Ecological Levin, S.A. and S.W.Pacala. 1997.Theories of simplifi- Genetics:The Interface. Springer-Verlag, . cation and scaling of spatially distributed processes. pp. Levin, S.A. 1978. Pattern formation in ecological 271-96. In: D.Tilman and P.Kareiva (eds.). Spatial Ecology: communities. pp. 433-65. In: J.H. Steele (ed.), Spatial The Role of Space in Population Dynamics and Interspecific Pattern in Plankton Communities. NATO Conference Series IV: Interactions. Princeton University Press, Princeton, NJ. Marine Sciences,Vol. 3 Plenum Press, NY. Levin, S.A., B.T. Grenfell,A. Hastings and A.S. Perelson. Levin, S.A. 1983. Some approaches to the modelling 1997. Mathematical and computational challenges in of coevolutionary interactions. pp. 21-65. In: M. Nitecki population biology and ecosystem science. Science. 275: (ed.), Coevolution. University of Chicago Press, Chicago, IL. 334-43.

10 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Levin, S.A., D. Cohen, and A. Hastings. 1984. Dispersal Okubo,A. and S.A. Levin (eds.) 2001. Diffusion and strategies in patchy environments.Theoretical Population Ecological Problems: Modern Perspectives, 2nd Edition. Biology 26(2): 165-91. Interdisciplinary Applied Mathematics,Vol 14. Springer, Levin, S.A., editor-in-chief. 2001. Encyclopedia of New York. pp. 467. Biodiversity,Volumes 1-5, Academic Press, San Diego. Paine, R.T. and S.A. Levin. 1981. Intertidal landscapes: Levin, S.A., H.C. Muller-Landau, R. Nathan, J. Chave. disturbance and the dynamics of pattern. Ecological 2003.The ecology and evolution of seed dispersal: Monographs 51(2): 145-78. A theoretical perspective. Annual Review of Ecology,Evolution, Plotkin, J. B. and S.A. Levin. 2001.The spatial and Systematics. 34: 575-604. distribution and abundances of species: Lessons from Levin, S.A., S. Barrett, S.Aniyar,W.Baumol, C. Bliss, tropical forests. Comments on Theoretical Biology 6: 251-78. B. Bolin, P.Dasgupta, P.Ehrlich, C. Folke, I.-M. Gren, Plotkin, J.B., J. Dushoff and S.A. Levin. 2002. C. S. Holling,A. Jansson, B.-O. Jansson, D. Martin, Hemagglutinin sequence clusters and the antigenic K.-G. Maler, C. Perrings and E. Sheshinsky.1998. evolution of influenza A virus. Proceedings of the National Resilience in natural and socioeconomic systems. Academy of Sciences,USA. 99(9): 6263-68. Environment and Development Economics 3: 225-36.

11 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Presentation address for the Dr A.H. Heineken Prize for Environmental Sciences Professor Johan Bouma

Delivered on the occasion of the presentation of the 2004 Heineken Prizes on October 1,2004

The jury of the 2004 Dr A.H. Heineken Prize for Environmental Sciences has unanimously decided to select you as the recipient of this award because of your highly significant, pio- neering research on Ecosystem Dynamics. Ecosystems are dynamic biological communities that inhibit specific areas and are continuously affected by internal and external processes in space and time.Your particular contribution involves the application of rigorous mathemati- cal complexity theory and analytical modeling to increase our basic understanding of the way in which ecosystems function at different spatial scales, ranging from the experimental plot to the landscape and beyond.You have shown that many of the properties of ecosystems vary according to fixed patterns, and that we can only understand what is going on at a particular spatial scale when we also understand what is happening at higher and lower scale levels.Your insights, which were generated by a fruitful combination of theory and empiricism, have led to fundamental changes in the discipline of ecology, which until the early nineties was strongly subdivided into various, rather independent subdisciplines.Your article, The Problem of Pattern and Scale in Ecology, became the most cited work in this field in the nineties. You are an exceptionally active and productive scientist.Your CV runs to dozens of pages and only lists significant achievements, no trivia.Your many publications include the 4800-page Encyclopedia of Biodiversity, of which you are Editor-in-Chief. Your qualities have been widely recognized. For example, you became a member of the American National Academy of Sciences in the year 2000, and you were presented with the Akira Okubo Lifetime Achievement Award in 2001.You are widely admired as a gifted speaker and excel- lent teacher with the rare ability to make your insights comprehensible to the general public. You also influence the international research agenda, for example as the chair of the Executive Committee of the International Institute for Applied Systems Analysis (IIASA), an inter- disciplinary research institute based in Austria, which studies the human dimensions of global change. A world that committed itself in Johannesburg in 2002 to sustainable develop- ment cannot achieve such worthy goals by good intentions alone. Research is needed to unravel the basic mechanisms of ecosystem dynamics, so as to allow the kind of understand- ing that is essential when judging actual conditions and when planning future developments. But there is more to consider than Ecosystem Dynamics by itself. can only be achieved when society at large is willing to embrace the challenges for the future,

12 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 including the major economic and social factors that must also be considered when inves- tigating ecosystems.Your ability to connect with scientists of other disciplines, for example economics, evolutionary biology and immunology,is very important in this context and pro- vides a clear channel to the practical study of biodiversity and nature conservation that are so important to society.You have in this way contributed to improving our understanding of the relationship between man and his environment, an important criteria for the Heineken Prize. Some prizes have the character of a career award, serving as the final recognition of individual achievement.That is certainly not the case here.You are highly active and intend to remain so.Your work will not only go on making a significant contribution to science, but it will also, in particular, provide support for all those who are struggling to put the some- what hazy concept of a sustainable world into practice.We hope to continue benefiting from your wisdom in the years ahead. The Academy also organized the 2004 Heineken Lectures. Four laureates were asked to lecture on their work to a broad audience at different locations. Unfortunately Jacques Le Goff was unable to travel to the Netherlands to give his Heineken Lecture, and his health did not permit him to come to Amsterdam for the Awards Ceremony. Instead, the Academy organized a symposium about Le Goff´s scholarly work. Mr Le Goff did, however, contribute a text about his recent work as an , which appears in this publication.

13 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 The Ecology of Complexity and the Complexity of Ecology

Dr A.H. Heineken Prize for Environmental Sciences 2004 Simon A.Levin

Professor Simon Levin received the Dr A.H.Heineken Prize for Environmental Sciences 2004 for his insights into the effects of scale on ecosystems.

14 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Our environment is at risk. We are facing a suite of global environmental problems, largely anthropogenic, of a magnitude without precedent since the last great extinctions. Principal among these are biodiversity loss, global atmospheric change, the depletion of critical resources on which human life depends, and the emergence and reemergence of devastating infectious diseases. How have we arrived at this fragile point in ’s history,and what can we do in response to mitigate our destructive impact? There is long history of research in resource management, especially of marine fisheries4. The mathematical theory is elegant, and traces back to Vito Volterra, the brilliant Italian mathematician who was elected President of Accademia Nazionale dei Lincei in 1923. Volterra, better known among mathematicians for his theoretical work, especially research on functional analysis and the theory of integral equations was attracted by his son-in-law, Umberto d’Ancona, to the problem of explaining the oscillations of the fisheries of the Adriatic. Volterra had had a long interest in applying mathematics to biology, as evidenced by a lecture he gave in 1900 at a convocation for the beginning of the academic year at the University of Rome. D’Ancona was himself a distinguished fisheries biologist, holding the Chair at Padua, and had accumulated statistics for the fluctuations of the Adriatic fish pop- ulations. D’Ancona’s conjecture was that predator-prey interactions drove the oscillations, and challenged Volterra to demonstrate this mathematically.For Volterra, this was an opportu- nity to put his theoretical interests to work, and he was up to the challenge. With simple mathematical models, he demonstrated how the predator-prey interaction could lead to oscillations, and went on to expand his work to address a broader range of ecological inter- actions1. The mathematical theory inspired by Volterra’s work and the parallel theoretical investigations of Alfred Lotka (1925) continues to this day,but has in general been a relative- ly sterile exploration of mathematical refinements, none with the insights of the masters. Meanwhile, fisheries management has been less than an unqualified success, and fisheries throughout the world are in collapse2. Why, in the face of such elegant theory,has fisheries management not been more successful? Marine ecosystems are complex systems, characterized by trophic interactions, nonlinearities and the potential for sudden flips. They are furthermore complex adaptive systems3 integrating phenomena at multiple scales. Despite Volterra’s contributions, marine fisheries have been treated largely as if they could be isolated from the ecosystems of which they are integral parts. Recently, there has been an increasing recognition of the essential nature of a whole-systems approach to marine fisheries4, but little has changed in terms of putting such principles into practice. The theory of infectious diseases similarly has a long and rich history,also going back about a century.The British bacteriologist, Sir Ronald Ross, won the for or Medicine in 1902 for his work on ; Ross developed perhaps the first dynamic models for the spread of an infectious disease5.The mathematical theory of infec- tious diseases has been a gem of the mathematical biology literature6, and has led to great advances in vaccination and other control strategies. Nonetheless, despite such elegant theo- ry, novel diseases continue to emerge, and old scourges reemerge. The problem here too is that these systems are complex systems, characterized by nonlinearities and the potential for

15 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 sudden flips. Indeed, they are also complex adaptive systems, integrating phenomena at mul- tiple scales.Treating diseases one at a time, without consideration of the broader-range inter- actions, ignores such complications as pathogen evolution and the emergence of bacterial antibiotic resistance. Disease agents such as influenza A have persisted for centuries as major scourges, despite the fact that a particular strain can infect an individual only once, because they change, adapting to the changing immune profile of the population. In disease systems, as in ecosystems and the biosphere more generally,macroscopic patterns emerge from evolu- tionary forces at much lower levels of organization; this is the essence of the challenge of complex adaptive systems. Not only are the natural systems we seek to manage complex adaptive systems; so, too, are the social and economic systems within which management must operate.They are also characterized by nonlinearities and the potential for sudden flips, and they too are driven largely by the actions of individual agents. Agreements such as the Kyoto accords are difficult to implement because we live in a global commons7 in which individual agents (including nations) act in their own self-interest, and in which the economic systems do not adequately incorporate the social costs. It is little wonder then that the collective actions of selfish agents can lead to global patterns of environmental degradation. The fundamental problems are hence ones of scale – spatial, temporal and organizational. To address these issues adequately, we must learn to extrapolate from small scales to large, from short time scales to long, and from leaves and cells to individuals, collectives, ecosystems and the bios- phere. The central theoretical question is to understand how ecosystems and the biosphere are organized; the central applied question is to learn what this understanding can tell us about effective management strategies. One of the most intriguing challenges for any natural scientist is to explain the regularities we see at the level of ecosystems and the biosphere, and the remarkable home- ostasis of conditions essential to life on the . , one of the world’s great- est Earth scientists, won the 1990 Heineken Prize largely for Gaia8, his presentation of ‘the earth as a living , in which there is a unity between animate and inanimate matter’. Few would question that the biota helps regulate aspects of the environment crucial to their survival; indeed, much of the concern about the current high rates of bio- diversity loss is because of the acknowledged role of biodiversity in mediating climate, and moderating environmental conditions more generally.The difficulty with for an evolutionary biologist, and indeed increasingly for Lovelock, is that whereas Gaia describes macroscopic regularities, evolution operates at much lower levels of organization, and not for the benefit of the whole system9. Lovelock (2002) wrote: ‘This idea was so contradictory to the views of evolutionary biologists that it was not long before , and other biologists challenged it.They pointed out that global regulation by the could never have evolved,because the organism itself was the unit of selection,not the Earth.In time I found myself agreeing with them.They were right:there was no way for organisms by themselves to evolve so that they could regulate the global environment.’

16 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004

It thus remains to be shown how such regularities in global temperature, and atmospheric , could have emerged over evolutionary and geological time, and become sustained over ecological time, given that and the other forces of evolution are strongest at those lower levels. Lovelock has recognized this, and asks further ‘I wondered, could the whole system, organisms and environment together, evolve self-regu- lation?,’ and applauded the efforts of Hamilton and Lenton (1998) to find common ground. In principle, the ideas are not so different from recent efforts at ‘niche construction’10; indeed, the differences are again issues of scale. Most evolutionary biologists recognize that organ- isms co evolve in some sense with their environments; the question is how those largely local results interact with the global scales of interest to Lovelock, and indeed ultimately to all of us. Evolution is not a process restricted to very long time scales. We have only to think of the rapid evolution of bacterial resistance, or heavy-metal tolerance in plants, to realize this. Even in the case of fisheries, the influence of humans in general, and harvest- ing strategies in particular, has led to evolutionary change in the life histories of key species, over relatively short periods of time. It is only in understanding the interplay among processes at these diverse scales that we can hope to gain a foothold in assuring a sustainable future for humanity. Ecosystems and the biosphere, as already discussed, are complex adaptive sys- tems; that is11, they are heterogeneous collections of individual agents that interact locally, and that evolve in their behaviors, genetics or distributions based on the outcomes of those interactions. Social and economic systems, physical and chemical systems, and the moral interconnections of society, also represent complex adaptive systems; and the common features of these interactive systems has led to an explosion of interest among diverse investi- gators12.The attractiveness of looking for common features of such systems is not new, and has found expression under the rubrics of pattern formation, collective phenomena, syner- getics13, and consilience14. As the interest has aged, however, it has also matured, integrating observation and mathematical theory,blending physical and social sciences with biology and evolutionary theory, and deepening our understanding of the interconnectedness of Earth’s biotic and a biotic components. In all of these systems, patterns emerge, to large extent, from phenomena at relatively low levels of organization – cells and individuals, small spatial scales and short temporal scales – but then those emergent patterns feed back, over larger and longer scales, to constrain the behavior of the lower-level dynamics. For the crucial task of understanding this interplay, it is powerful to develop agent-based models, conceptual or mathematical, to explore the extent to which observed macroscopic patterns self-organize from these, and the extent to which those patterns are imposed by external forcing. In this lecture, I will follow this approach in considering the fundamental problem of consumption in biological communities, and what we can learn from this for understanding the causes and remedies for over consumption in human societies15. Thus, I will divide my discussion into three parts: • the evolutionary ecology of consumption • the evolutionary ecology of cooperation and reduced consumption • the social norms and consumption in human societies.

18 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 The evolutionary ecology of consumption I will not attempt a complete discussion of theories of the evolution of consumption in natu- ral communities; the ecological literature is voluminous on this theme. Noteworthy among recent work is the monograph of Tilman (1982), and the central point that among species competing for a single limiting resource, only one will survive – that which can maintain itself at the lowest level of the resource. This result is an expansion of the classical theory of competitive exclusion16; but Tilman uses it to build the foundations of a theory of compe- tition for multiple resources. Even more recent has been the development of a stoichiometric theory of resource utilization17, which provides a framework for helping to address some long-standing problems in marine ecology, with implications for resolving the challenges of Gaia. Alfred Clarence Redfield, one of the first staff members recruited by Henry Bryant Bigelow to the Woods Hole Oceanographic Institution, discovered that the average atomic proportions of C, N and P in marine were approximately in the ratios 105:15:1, virtually identical to the biogenic ratios in the open ocean. Is this the result of organisms simply having evolved to match the ambient conditions, somehow pegged at their values independently of the biota; or does this regularity reflect biotic control over their environ- ment, a la Gaia, at just the right conditions for their survival? The answer remains open, although Redfield believed that bacterial nitrogen fixation and denitrification played an essen- tial bottom-up role in the control process. Clearly, what is needed are theoretical approaches that cross scales, integrating the ecology and evolution of nutrient utilization, combining approaches such as those of Tilman and of Sterner and Elser with evolutionary models, at least at the phenotypic level. In investigating this question, the challenges are to determine to what extent the ratios in the water column are simply reflecting those in the biota, and to what extent the reverse is true.What determines phytoplankton stoichiometry, and what determines alloca- tion to uptake versus growth? Ecological models, incorporating well-established uptake rela- tions and growth functions can describe the interactions of species of known stoichiometry with their resources, and competition among types.Then, on evolutionary time scales, muta- tion and other mechanisms can generate variation, allowing examination of natural selec- tion. Led by postdoctoral fellows Chris Klausmeier,Tanguy Daufresne and Elena Litchman, we have been adopting this approach with great success, providing strong support for the thesis that the biota indeed are controlling the ratios, and that it is the stoichiometry of and ribosome’s that determines the evolutionary outcome18. Coupling this approach with realis- tic models of the fluid dynamics of the oceans (together with fluid dynamicist Glenn Flierl), and the superposition of nutrient pulses, we have high hopes of explaining the community ecology of phytoplankton, and the dynamics of coexistence of multiple stoichiometries. The focus on the importance of fluid dynamics directs attention as well to the role of mixing. In general, there is considerable evidence19 that the localization of interactions in space is a major factor allowing coexistence of species, and that increased homogenization reduces biodiversity. Terrestrial systems also show characteristic Redfield ratios, but they are in general very different than are the marine ratios20. Furthermore, nitrogen fixation is often

19 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 lacking even in environments in which nitrogen is limiting21; and, more generally,organisms simply do not make use of all the resources that they might.Why? Could the reduced levels of mixing in terrestrial systems contribute to such prudence; and if so, how? We (Zea et al. in prep) have mimicked the ecological-evolutionary approach we used for marine systems in terrestrial systems, considering water-use patterns among plants in arid environments, and varying the level of mixing of the water tables.As mixing is increased in the model, evolution drives plants to become more selfish, increasing water utilization.This result complements previous theoretical studies by a variety of authors22 in other systems, and is an evolutionary manifestation of Hardin’s Tragedy of the Commons: There is less incentive to save for the future when savings must be shared with others.This finding, both depressing and intuitive, provides a natural segue into the second part of this disquisition.

The evolutionary ecology of cooperation and reduced consumption Altruism was a puzzle even for Charles Darwin. If natural selection is about the survival of the fittest, how are obvious examples of altruism sustained? Why do the females of the haplodiploid insects sacrifice their own reproductive fitness to help their sisters? The great evolutionary biologist J.B.S. Haldane (1948) summarized the essential truth in his compact statement that he would give up his life for two brothers, or for eight cousins, reflecting the fact that he shared 1/2 of his genes with each brother, and 1/8 with each cousin. It was left to W.D.Hamilton to work out the theory more completely23, developing the notion of extend- ed fitness. In the haplodiploid insects, the males arise from unfertilized eggs, meaning that two full sisters share 3/4 of their genes.Thus, true eusociality can evolve much more easily in the haplodiploid species than in fully diploid ones, because apparently altruistic individuals are simply finding a better way of getting their genes into the next generation. Extensions of this idea led to the controversial notion of the selfish gene24. Tight genetic relatedness is only one way to tighten the reward loop to the altru- ist; repeated interactions with the same individuals in space or time can achieve the same result. Thus, models with reduced mixing, which hence assume that individuals interact more often with their close neighbors, also lead more easily to the evolution of altruism and cooperation25. Unfortunately, tight feedback loops do not guarantee cooperative behavior; they can just as easily lead to greed and monopolistic domination of resources, and to mech- anisms such as allelopathy for interference with others26. In human societies, social norms have evolved to counteract this drift towards selfishness, but again they can also lead to self- ishness at the inter-group level. Indeed, as cultures come more into contact and conflict at broader scales, it is the absence of more widely agreed-upon social norms that inhibits the peaceful resolution of conflicts. In the last section, I turn to these problems, and what evolu- tionary biology might tell us about their resolution.

Social norms and consumption in human societies How can we apply the lessons of studying the evolution of consumption to patterns of con- sumption, and over consumption27, in human societies? The answer lies in part in realizing

21 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 that those patterns are to large extent governed by social norms, and thereby sustained because of the robustness of common practice. In 1997, the U.S.public broadcasting network PBS aired the program ‘Affluenza,’ by which they mean ‘an epidemic of stress, waste, over consumption and environmental decay’28. Consumptive patterns are in a sense social diseases, spread by infectious contact, and sustained by common practice. Indeed, as with other social norms that became widespread, their popularity subsequently can lead to governmental measures to sustain and strengthen such practices. To create and foster multil-level social organizations and governance, we need not only to develop global organizations, but also to foster local level organizations that confer robustness on sustainable management29. Efforts to replace traditional bottom-up governance with top-down efforts that seek to impose uniformity can have disastrous consequences; no example is clearer than that of the water- temples of Bali, where government intervention to try to improve a self-organized form of governance led to collapse30. Fortunately, the collapse was reversible once the traditional system was restored. Lansing’s analysis of this example brilliantly illustrates the perils and potential of management in a complex adaptive ecological-social system. As discussed earlier, cooperation arises most easily when interactions are tight. Closely related individuals, or those that interact frequently, form cooperative groups that increase their ability to compete with other groups. Such groups are held together by com- mon practice, and often by rituals that bind individuals to their groups. As group influence spreads, group size increases, often by higher-order levels of cooperation, as the group them- selves become the basic units of interaction. Larger groups are more influential, but are like- ly to be more heterogeneous. Individuals in large groups benefit from the increased power of the group, but suffer from a reduced ability to influence group dynamics.Thus the formation and dissolution of groups is a complex process involving fitness trade-offs, and the spread of associated behaviors. It is clear that in the dynamics of these social processes lie the keys to effective environmental management. A great challenge before us is thus to understand the dynamics of social norms31 – how they arise, how they spread, how they are sustained and how they change. Models of these dynamics have many of the same features as models of epidemic spread32 – no great sur- prise, since many aspects of culture have the characteristics of being social diseases. 1998 Heineken award winner Paul Ehrlich and I have been directing our collective energies to this problem, convinced that it is as important to understand the dynamics of the social systems in which we live as it is to understand the ecological systems themselves. Understanding the links between individual behavior and societal consequences, and characterizing the networks of interaction and influence, create the potential to change the reward structures so that the social costs of individual actions are brought down to the level of individual payoffs. It is a daunting task, both because of the amount we still must learn, and because of the ethical dilemmas that are implicit in any form of social engineering. But it is a task from which we cannot shrink, lest we squander the last of our diminishing resources.

22 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Acknowledgments I gratefully acknowledge helpful comments by Paul Ehrlich,Lars Hedin,Ann Kinzig, Carole Levin and Elinor Ostrom,and the invaluable assistance of Amy Bordvik.

Literature cited 1 Volterra 1926 2 Myers & Worm 2003; Pauly et al. 1998, 2002 3 Levin 1998, 1999 4 Policansky et al. 1999; Duda & Sherman 2002 5 Ross 1911 6 Anderson & May 1979; May & Anderson 1979; Levin et al. 1989 7 Hardin 1968; Levin 1999 8 Lovelock 1979 9 Lovelock 1983; Ehrlich 1991; Kirchner 1991; Schneider & Boston 1991; Levin 1999 10 Odling-Smee et al. 2003 11 Levin 1998, 1999 12 Gell-Mann 1994; Holland 1995 13 Haken 1983 14 Wilson 1998 15 Arrow et al. in press 16 see Levin 1970 17 Sterner & Elser 2002 18 Klausmeier et al. 2004 a,b 19 Levin & Paine 1974; Levin 1976 20 Hedin 2004; McGroddy et al. in press 21 Vitousek et al. 2002 22 Kinzig & Harte 1998; Klopfer 1997 23 Hamilton 1964 24 Dawkins 1976 25 Nowak & May 1992; Durrett & Levin 1994; Miller 1996 26 Chao & Levin 1981; Durrett & Levin 1997; Kerr et al. 2002 27 Arrow et al. in press 28 http://www.pbs.org/kcts/affluenza/escape 29 Dietz et al. 2003 30 Lansing 1993 31 Axelrod 1986 32 Durrett & Levin, in press

23 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 References Evolution 10: 1-16. Anderson, R.M., and R.M. May.1979. Population biology Hardin, G. 1968.The tragedy of the commons. of infectious diseases: Part I. Nature 280: 361-367. Science 162: 1243-1248. Arrow,K., G. Daily,P.Dasgupta, P.Ehrlich, L. Goulder, Hedin, L.O. 2004. Global organization of terrestrial G. Heal, S. Levin, K.-G. Maler, S. Schneider, D. Starrett, plant – nutrient interactions. Proceedings of the National and B.Walker. 2004.Are we consuming too much? Academy of Science USA 101: 10849-10850. Journal of Economic Perspectives. In press. Holland, J. 1995. Hidden Order.How Adaptation Builds Axelrod, R. 1986.An evolutionary approach to norms. Complexity. Addison Wesley,Reading, MA. American Political Science Review 80: 1095-1111. Kerr, B., M.A.Riley,M.W.Feldman, and B.J.M. Bohannan. Chao, L., and B.R. Levin. 1981. Structured habitats 2002. Local dispersal promotes biodiversity in a real-life and the evolution of anticompetitor toxins in bacteria. game of rock-paper-scissors. Nature 418: 171-174. Proceedings of the National Academy of Sciences,USA 78: Kinzig,A.P.,and J. Harte. 1998. Selection of 6324-6328. microorganisms in a spatially explicit environment and Dawkins, R. 1976. The Selfish Gene. Oxford University implications for plant access to nitrogen. The Journal of Press, New York. Ecology 86: 841-853. Dietz,T., E. Ostrom and P.C. Stern 2003,The Struggle Kirchner, J.W.1991.The Gaia hypothesis:Are they to Govern the Commons, Science 302: 1907-1912. testable? Are they useful? Pages 38-46 in S. H. Schneider Duda,A.M., and K. Sherman. 2002.A new imperative and P.J. Boston, eds. Scientists on Gaia. MIT Press, for improving management of large marine ecosystems. Cambridge, MA. Ocean & Coastal Management 45: 797-833. Klausmeier, C.A., E. Litchman,T. Daufresne, and S.A. Durrett, R., and S.A. Levin. 1994.The importance of Levin. 2004a. Optimal nitrogen-to-phosphorus being discrete and (spatial). Theoretical Population Biology 46: stoichiometry of phytoplankton. Nature 429: 171-174. 363-394. Klausmeier, C.A., E. Litchman, and S.A. Levin. 2004b. Durrett, R., and S.A. Levin 1997.Allelopathy in spatially Phytoplankton growth and stoichiometry under multiple distributed populations. Journal of Theoretical Biology 185: nutrient limitation. Limnology and Oceanography 49: 1463- 165-171. 1470. Durrett, R. and S.A. Levin 2004. Can stable social Klopfer, E. 1997. Evolution of intermediate exploitation groups be maintained by homophilous imitation alone? rates in exploiter-victim systems. Ph.D. dissertation, Journal of Economic Behavior and Organization. In press. University of Wisconsin, Madison. Ehrlich, P.1991. Co evolution and its applicability to Lansing, J.S. 1993. Priests and Programmers:Technologies of the Gaia hypothesis. Pages 19-22 in S. H. Schneider and P.J. Power in the Engineered Landscape of Bali. Princeton University Boston, eds. Scientists on Gaia. MIT Press, Cambridge, MA. Press, Princeton, NJ. Gell-Mann, M. 1994. The Quark and the Jaguar:Adventures Levin, S.A. 1970. Community equilibria and stability, in the Simple and Complex.W.H.Freeman and Co., New York. and an extension of the competitive exclusion principle. Haken, H. 1983. Synergetics. Springer, New York. American Naturalist 104: 413-423. Haldane, J.B.S. 1948.The theory of cline. Journal of Levin, S.A. 1976. Population dynamic models in Genetics 48: 227-284. heterogeneous environments. Annual Review of Ecology Hamilton,W.D. 1964.The genetical evolution of social and Systematics 7: 287-311. behavior (I and II). Journal of Theoretical Biology 7: 1-16; Levin, S.A. 1998. Ecosystems and the biosphere 17-32. as complex adaptive systems. Ecosystems 1: 431-436. Hamilton,W.D., and T.M. Lenton. 1998. Spora and Gaia: Levin, S.A. 1999. Fragile Dominion:Complexity and How microbes fly with their clouds. Ethology Ecology & the Commons. Perseus Books, Reading, MA.

24 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Levin, S.A.,T.G. Hallam, and L. J. Gross, eds. 1989. Applied 1999. Sustaining Marine Fisheries. National Academy Press, Mathematical Ecology.Biomathematics 18. Springer-Verlag, Washington, DC. Heidelberg. Ross, R. 1911. Some quantitative studies in Levin, S.A., and R.T. Paine. 1974. Disturbance, patch epidemiology. Nature 87: 466-467. formation, and community structure. Proceedings of the Schneider, S.H., and P.J. Boston, Eds. 1991. Scientists National Academy of Sciences USA 74: 2744-2747. on Gaia. MIT Press, Cambridge, MA. Lotka,A.J. 1925. Elements of Physical Biology.Williams Sterner, R.W.,and J.J. Elser. 2002. Ecological Stoichiometry. and Wilkins, Baltimore, MD. Princeton University Press, Princeton, NJ. Lovelock, J.E. 1979. Gaia:A New Look at Life on Earth. Tilman, D. 1982. Resource Competition and Community , Oxford. Structure. Princeton University Press, Princeton, NJ. Lovelock, J.E. 1983. :A cybernetic proof of Vitousek, P.M.,K. Cassman, C. Cleveland,T. Crews, C.B. the Gaia hypothesis. Co evolution Quarterly 38: 66-72. Field, N.B. Grimm, R.W.Howarth, R. Marino, L. Martinelli, Lovelock,, J.E. 2002.What is Gaia? Resurgence. Issue 211. E.B. Rastetter, and J.I. Sprent. 2002.Towards an http://resurgence.gn.apc.org/issues/lovelock211.htm ecological understanding of biological nitrogen fixation. May,R.M., and R.M.Anderson. 1979. Population 57/58: 1-45. biology of infectious diseases: Part II. Nature 280: 455-461. Volterra,V.1926.Variazioni e fluttuazioni del numero McGroddy,M.E.,T. Daufresne, and L.O. Hedin. 2004. d’individui in specie animale conviventi. Mem R Accad Scaling of C:N:P stoichiometry in forest ecosystems Nazionale del Lincei (Ser. 6) 2: 31-113. worldwide: Implications of terrestrial Redfield-type ratios. Wilson, E.O. 1998. Consilience:The Unity of Knowledge. Ecology. In press. Vintage Books, New York. Miller, J.H. 1996.The co evolution of automata in the repeated prisoner’s dilemma. Journal of Economic Behavior and Organization 29: 87-112. Myers, R.A., and B.Worm. 2003. Rapid worldwide depletion of predatory fish communities. Nature 423: 280-283. Nowak, M.A., and R.M. May.1992. Evolutionary games and spatial chaos. Nature 359: 826-829. Odling-Smee, F.J., K.N. Laland, and M.W.Feldman. 2003. Niche Construction:The Neglected Process in Evolution. Princeton University Press, Princeton, NJ. Pauly,D.,V.Christensen, J. Dalsgaard, R. Froese, and J.Torres, Francisco. 1998. Fishing down marine food webs. Science 279: 860-863. Pauly,D.,V.Christensen, S. Guénette,T.J. Pitcher, U.R. Sumaila, C.J.Walters, R.Watson, and D. Zeller. 2002. Toward in world fisheries. Nature 418: 689-695. Policansky,D., H. Mooney,D.L.Alverson, H. Bingham, J. Clark, F. Grassle, E. Hofmann, E. Houde, S. Levin, J. Lubchenco, J. Magnuson, B. McCay,G. Munro, R. Paine, S. Palumbi, D. Pauly,E. Pikitch,T. Powell, and M. Sissenwine

25 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Heineken Lecture Program 2004

The Heineken Lectures were presented on 28 September and 30 September 2004.

Simon Levin laureate of the Dr A.H. Heineken Prize for Environmental Sciences 2004 Heineken Lecture The Ecology of Complexity,and the Complexity of Ecology Royal Tropical Institute,Amsterdam

Daan van Golden laureate of the Dr A.H. Heineken Prize for Art 2004 Heineken Lecture Red Or Blue,Some Words Of Artful Wisdom De Ateliers,Amsterdam

Jacques Le Goff laureate of the Dr A.H. Heineken Prize for History 2004 Symposium The Other De Balie,Amsterdam

Andrew Fire laureate of the Dr H.P.Heineken Prize for Biochemistry and Biophysics 2004 Heineken Lecture How Cells Respond to Genetic Change , Utrecht

Elizabeth Blackburn laureate of the Dr A.H. Heineken Prize for Medicine 2004 Heineken Lecture Telomeres and Telomerase in Health and Disease Utrecht University,Utrecht

26 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Audience and publicity for the Heineken Lectures in 2004

The Heineken Lectures are intended for a broad audience. Students, scientists, Academy members, but also laymen who are interested in the field of study or the research associated with one or more of the Heineken Prizes can attend the Heineken Lectures free of charge.

In previous years, the laureates gave their Heineken Lectures during the course of a single Academy session at the Trippenhuis Building in Amsterdam, the headquarters of the Royal Netherlands Academy of Arts and Sciences. Starting in 2002, the Heineken Lectures were given at different locations throughout the Netherlands in order to reach a broader audience. In 2004, the Heineken Lectures were not only delivered at different locations, but also on different dates, drawing more people than ever before. More than seven hundred people attended one or more of the Heineken Lectures.

The large number of attendees is partly the result of a major campaign launched in 2004 to generate more publicity for the Heineken Prizes, and in particular for the Heineken Lectures. The campaign, run by the Royal Netherlands Academy of Arts and Sciences and Heineken International, consisted of leaflets, announcement posters, free tickets, the website Heinekenprizes.org, a special issue of the Academy’s quarterly magazine Akademie Nieuws, and a booklet with more information about the Heineken Prizes and the laureates in 2004.

Between April and October of 2004, the Heineken Prizes website of the Royal Netherlands Academy of Arts and Sciences, www.knaw.nl/heinekenprizes, provided updated information on the 2004 Heineken Prizes.The site now offers a detailed review of the event, with infor- mation on the background and organization of the prizes, the nomination procedure, and the laureates, as well as press information (including photos and documentation).

27 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Secondary School Project about the Heineken laureates

In 2004, the Royal Netherlands Academy of Arts and Sciences initiated a secondary school project on the 2004 Heineken Prizes on Kennisnet, the Internet organization for primary, secondary and vocational education in the Netherlands.

The Royal Academy hired a professional teaching organization to develop five kits that help secondary school students write papers on the work and research of the five laureates of the 2004 Heineken Prizes.The kits cover the fields of Biochemistry and Biophysics (RNA-inter- ference), Medicine (telomerase), Environmental Sciences (ecological systems) and History (the way an average person in the Middle Ages looks upon the world around him).The fifth kit is about the life and work of Dutch artist Daan van Golden.The information provided in the kits was written by Dutch university students enrolled in a variety of different programs.

By offering secondary school students kits like these, the Royal Academy is helping to acquaint them with top scientists and top scientific research.The hope is that they will then have a better idea of what they would like to study after graduation. From October to December 2004, almost three thousand students, teachers and other people inspected the Academy’s kits.

The five kits can be found on the website www.werkstuknetwerk.nl.

28 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 General information

The Heineken Prizes:five prizes for outstanding contributions to the arts and sciences

Every two years the Dr H.P. Heineken Foundation and the Alfred Heineken Fondsen Foundation award four prizes – a cash gift of 150.000 USD and a crystal symbol – to scien- tists in the disciplines of Biochemistry and Biophysics, Medicine, Environmental Sciences and History for outstanding contributions to their field of study and one prize for the performing arts to a Dutch artist (50.000 EUR).

The selection of the winners for the Heineken Prizes has been entrusted to the Royal Netherlands Academy of Arts and Sciences. The Academy’s Arts and Sciences Divisions have appointed special committees to carry out this task.The jury of the Dr A.H. Heineken Prize for Art consists of three members of the Academy complemented by experts in the particular artistic field.

The Academy also organized the 2004 Heineken Lectures. Four laureates were asked to lecture on their work to a broad audience at different locations.

29 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 List of Heineken laureates

Dr H.P.Heineken Prize for Biochemistry and Biophysics

1964 1967 Jean L.A. Brachet 1970 1973 1976 Laurens L.M. van Deenen 1979 1982 1985 Bela Julesz/Werner E. Reichardt 1988 Thomas R. Cech 1990 1992 1994 Michael J. Berridge 1996 Paul M. Nurse 1998 Tony J. Pawson 2000 James E. Rothman 2002 Roger Y.Tsien 2004 Andrew Z. Fire

Dr A.H. Heineken Prize for Art

1988 1990 1992 1994 Matthijs Röling 1996 1998 2000 2002 2004 Daan van Golden

30 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Dr A.H. Heineken Prize for Medicine

1989 Paul C. Lauterbur 1990 Johannes J. van Rood 1992 1994 1996 1998 Barry J. Marshall 2000 Eric R. Kandel 2002 Dennis J. Selkoe 2004 Elizabeth H. Blackburn

Dr A.H. Heineken Prize for History

1990 1992 1994 Peter R.L. Brown 1996 Heiko A. Oberman 1998 Mona Ozouf 2000 2002 2004 Jacques Le Goff

Dr A.H. Heineken Prize for Environmental Sciences

1990 James E. Lovelock 1992 Marko Branica 1994 BirdLife International (Colin J. Bibby) 1996 Herman E. Daly 1998 Paul R. Ehrlich 2000 Poul Harremoës 2002 Lonnie G.Thompson 2004 Simon A. Levin

31 THE DR A.H. HEINEKEN PRIZE FOR ENVIRONMENTAL SCIENCES 2004 Colophon

©2005 Royal Netherlands Academy of Arts and Sciences No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher.

P.O. Box 19121, 1000 GC Amsterdam, the Netherlands Telephone: +31(0)20-5510700 Fax: +31(0)20-6204941 E-mail: [email protected] www.knaw.nl

ISBN 90-6984-437-0

The Heineken Prizes website, www.knaw.nl/heinekenprizes, has more information on the background and organization of the Heineken Prizes, the nomination procedure, the laureates and their research. The site also provides press information (including photos and documentation).

Editing

Royal Netherlands Academy of Arts and Sciences Photography

Christiaan Krouwels Graphic design

Design Studio Hans Kentie BNO Colour separations

United Graphics Printing

Boom Planeta Graphics Binding

J.Voetelink & Zoon bv The Ecology of Complexity and the Complexity of Ecology Ecosystems and the biosphere are complex adaptive systems in which macroscopic features, such as robustness, emerge largely from processes at lower scales of organization. A fundamental challenge, then, is to relate processes and patterns occurring at diverse scales of space, time and organizational complexity. What maintains the macroscopic patterns of the biosphere, such as the distributions and abundances of species? How do these regularities emerge at the level of ecosystems and the biosphere, given that evolution there is mediated largely at much smaller scales of organization? How are the responses and actions of individual plants and animals translated into phenomena at higher levels of organization, and how do these phenomena feed back to influence the fates of individuals? How do complex networks of interaction arise, and what are the implications for the robustness of these systems in the face of anthropogenic disturbance? Professor Simon Levin’s lecture explores these issues and the lessons the evolution of natural systems can teach us about cooperation in human societies, the development of social norms and institutions, and the management of resources in a global commons. T h e E c o l o g y o f C o m p l e x i t y a n d t h e C o m p l e x i t y o f E c o l o g y