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    http://greencraft.co.uk/

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    farm3.static.ickr.com

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    Can we grow economically without

    compromising options for future generations

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    (Brundtland report)

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    Developing a comprehensive

    framework for answering thatquestion is the rst order of business

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    Measurement of utility Aggregation and intertemporal social welfare Are we consuming too much?

    Implications for policy

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    The challenges are interdisciplinary

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    www.colorado.edu

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    JSTOR: The Journal of Economic Perspectives, Vol. 18, No. 3... http://www.jstor.org/stable/3216811

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    Sustainability means many things

    www.waikato.govt.nz/enviroinfo

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    Sustainability means many things

    www.serconline.org

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    Are the services we derive fromecosystems sustainable?

    marinebio.org/i/

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    Key research questions What are services?

    Direct, indirect, aesthetic/ethical

    and how do they depend upon features of ecosystems?

    Connections across scales

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    Characteristic regularities in

    macroscopic patterns exist in allecosystems

    www.yale.edu/yibs

    www.csiro.au

    www.bio.unc.edu

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    There are striking regularities in such macroscopicpatterns, independent of much microscopic detail

    Volkov, Banavar, Hubbell and MaritanNature 424 , 1035-1037

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    This implies a need to relatephenomena across scales, from

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    Sustainability must focus on macroscopicfeatures, while recognizing that control of those

    rests at lower levels of organization

    www.pitt.edu/~jdnorton

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    Studying emergence and evolution Agent-based and hierarchical models of self-

    organizing systems

    Statistical mechanics of ensembles of agents Emergent description of macroscopic dynamics

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    u and K from ECCO2 GCM Phyto growthRemineralization &other sources

    Growth Mortality GrazingSinking

    MJ Follows et al , Science 315 , 1843 (2007)

    Towards a Trait-Based Ecology, the MIT-DARWIN Model

    Wunsch & P Heimbach, Physica D 230 ,197 (2007)

    N/P/Z= nutrients/phytoplankton/zooplankton

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    Diatoms

    Prochlorococcus

    Synechococcus

    Large eukaryotes

    Follows, Dutkiewicz, Chisholm,

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    Key research questions What are services, and how do they depend upon

    features of ecosystems? Direct, indirect, aesthetic/ethical Connections across scales

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    Thus, macroscopic regularities of systems emerge fromecological and evolutionary interactions on ner scales

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    Key research questions What are services, and how do they depend upon

    features of ecosystems?

    What maintains characteristic features? Whatsustains robustness of key emergent properties of coupled human-environmental systems?

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    Ecosystems and the Biosphere are

    Complex Adaptive SystemsHeterogeneous collections of individual units(agents) that interact locally, and evolve

    based on the outcomes of those interactions.

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    Features of CAS Self-organization Multiple stable states, path dependence, hysteresis Contagious spread and systemic risk Multiple time scales, and potential for

    destabilization and regime shifts through slow-time-scale evolution

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    Characterizing the robustness/vulnerability of

    complex adaptive human-environment systems

    How do systems self-organize over time? What makes systems robust? Does robustness increase over time, or does

    system evolution carry the seeds of its owncollapse?

    What are the indicators of the erosion of robustness?

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    REVIEWS

    Early-warning signals for critical transitionsMarten Scheffer 1, Jordi Bascompte 2, William A. Brock 3, Victor Brovkin 5, Stephen R. Carpenter 4, Vasilis Dakos 1,Hermann Held 6, Egbert H. van Nes 1, Max Rietkerk 7 & George Sugihara 8

    Complex dynamicalsystems, rangingfrom ecosystems to financialmarketsand the climate, can havetippingpoints at whicha sudden shiftto a contrasting dynamicalregime may occur. Although predicting suchcritical points before theyare reached

    is extremely difficult, work in different scientific fields is now suggesting the existence of generic early-warning signals thatmay indicate for a wide class of systems if a critical threshold is approaching.

    It is becoming increasingly clear that many complex systems havecritical thresholdsso-calledtippingpointsatwhich thesystemshifts abruptly from one state to another. In medicine, we havespontaneous systemic failures suchas asthma attacks 1 or epileptic

    seizures 2,3; in global finance, there is concern about systemic marketcrashes 4,5; in the Earth system, abrupt shifts in ocean circulation orclimate mayoccur 6; andcatastrophic shiftsin rangelands, fishpopula-tions or wildlife populations may threaten ecosystemservices 7,8.

    It isnotablyhardto predict suchcriticaltransitions,becausethe stateof the system may show little change before the tipping point isreached. Also, models of complex systems are usually not accurateenough to predict reliably where critical thresholds may occur.Interestingly, though, it now appears that certain generic symptomsmayoccurin awideclassof systemsastheyapproacha critical point.Atfirstsight, itmayseemsurprisingthatdisparatephenomenasuch asthecollapse of an overharvested population and ancient climatic transi-tions couldbe indicatedby similar signals.However,as wewill explainhere, the dynamics of systems near a critical point havegeneric prop-erties,regardlessof differences inthe details ofeachsystem 9. Therefore,sharp transitions in a range of complex systems are in fact related. Inmodels, critical thresholds for such transitionscorrespond to bifurca-tions 10. Particularly relevant are catastrophic bifurcations (see Box1for an example), where, once a threshold is exceeded, a positive feed-back propels thesystem through a phaseofdirectional change towardsa contrasting state. Another important class of bifurcations are thosethat markthe transition froma stableequilibriumto a cyclicor chaotic

    r r F n m n l hif h r in m h n h

    considered to capture the essence of shifts at tipping points in a widerange of natural systems ranging from cell signalling pathways 14 toecosystems 7,15 and the climate 6. At fold bifurcation points ( F 1 and F 2,Box1), the dominant eigenvalue characterizing the rates of changearound the equilibrium becomes zero. This implies that as the systemapproaches such critical points, it becomes increasingly slow in re-covering from small perturbations (Fig. 1). It can be proven that thisphenomenon will occur in any continuous model approaching a foldbifurcation 12. Moreover, analysis of various models shows that such

    slowing down typically starts far from the bifurcation point, and thatrecovery rates decrease smoothly to zero as the critical point isapproached 16. Box 2 describes a simple example illustrating this.

    The most straightforward implication of critical slowing down isthat the recovery rate after small experimental perturbation can beused as anindicator of how close a systemis to a bifurcation point 16.Because it is the rate of change close to the equilibrium that matters,such perturbations may be very small, posing no risk of driving thesystem over the threshold. Also, models indicate that in spatially extensive systems at risk of systemic collapse, small-scale experi-mental probing may suffice to test the vicinity of the threshold forsuch a large-scale transition. For instance, it has been shown thatrecovery times after local perturbation increase in models of frag-mented populations approaching a thresholdfor global extinction 17.

    Formost naturalsystems, it would be impracticalor impossible tomonitor them by systematically testing recovery rates. However,almost all real systems are permanently subject to natural perturba-i n I n h n h if r i n i r h in h

    Vol 461 j3 September 2009 jdoi:10.1038/nature08227

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    2008

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    2008

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    2008

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    Key research questions What are services, and how do they depend upon

    features of ecosystems?

    What maintains those features? Management of human-environmental systems to

    sustain services

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    Rohde

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    But adequate action to address themhas been lacking

    www.edie.net

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    The central issues are issues of behavior and culture

    Intergenerational and intragenerational equity Public goods and common pool resources Cooperation in the Commons Social norms and institutions Role of leadership and collective behavior

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    Leadership and collective behavior

    Carere

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    Role of leadership and collectivedecision-making

    Couzin, Krause, Franks, Levin

    Couzin/BBC

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    g 1

    C o

    l l e c

    t i v e

    d e c i s

    i o n - m a

    k i n g

    Trend setter

    Copier

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    1 informed individuals in group of 100.

    C o

    l l e c

    t i v e

    d e c

    i s i o n - m a

    k i n g

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    10 informed individuals in group of 100.

    C o

    l l e c

    t i v e

    d e c

    i s i o n - m a

    k i n g

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    Animal groups may be led by a small

    number of individuals

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    So too are human societies

    42http://interactivedemocracy.blogspot.com/

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    Consensus may also be emergent

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    Uninformed individuals promote democratic consensus inanimal groups

    Iain D. Couzin 1, Christos C. Ioannou 1, Gven Demirel 2, Thilo Gross 2, Colin J. Torney 1, AndrewHartnett 1, Larissa Conradt 3, Simon A. Levin 1, Naomi E. Leonard 4

    1. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, Ne Jersey 08544, USA

    2. Max-Planck Institute for Physics of Complex Systems, Nthnitzer Str., 01187 Dresden, Germany

    3. John Maynard Smith Building, School of Life Sciences, University of Sussex, Falmer, Brighton, BN1

    9QG, UK

    4. Department of Mechanical and Aerospace Engineering, Princeton University, Princeton, New Jersey

    08544, USA

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    Furthermore, we discount The future

    www.elements4health.com

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    We discount The future The interests of others

    www.improvingyourworld.co

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    How much should we leave to future

    generations?

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    Intergenerational equity

    www.dot.ca.gov/hq

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    The problem of intergenerational transferof resourceshas strong parallels in evolutionary theory

    R.Klopfer

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    Intergenerational resource transfers with randomoffspring numbersKenneth J. Arrowa and Simon A. Levinb,1

    aDepartment of Economics, Stanford University, Stanford, CA 94305-6072; andbDepartment of Ecology and Evolutionary Biology, Princeton University,Princeton, NJ 08544-1003

    Contributed by Kenneth J. Arrow, May 26, 2009 (sent for review March 29, 2009)

    A problem common to biology and economics is the transfer ofresourcesfromparentsto children.We consider theissue under theassumption that the number of offspring is unknown and can berepresented as a random variable. There are 3 basic assumptions.The rst assumption is that a given body of resources can bedivided into consumption (yielding satisfaction) and transfer tochildren. The second assumption is that the parents welfareincludes a concern forthe welfare of their children;this is recursive

    in the sense that the childrens welfares include concern for theirchildren and so forth. However, the welfare of a child from a givenconsumptionis counted somewhat differently(generallyless) thanthat of theparent(thewelfare of a childis discounted). Thethirdassumption is that resources transferred may grow (or decline). Ineconomic language, investment, including that in education ornutrition, is productive. Under suitable restrictions, precise formu-las for the resulting allocation of resources are found, demonstrat-ing that, depending on the shape of the utility curve, uncertaintyregarding thenumber of offspringmay or maynot favor increasedconsumption. The results imply that wealth (stock of resources)will ultimately have a log-normal distribution.

    allocation intergenerational transfers life history theory uncertainty

    ping generations, offspring produced early in life are more valuable than those produced later because those offspring canalso begin reproduction earlier. This is analogous to the classicinvestment problem in economics, in that population growthimposes a discount rate that affects when one should haveoffspring. The flip side is that early reproduction compromisesthe parents ability to care for its children, and that increasednumber of offspring reduces the investment that can be made in

    each. Again, thebest solutiongenerallyinvolves compromise andan intermediate optimum. A particularly clear manifestation of this tradeoff involves the

    problem of clutch or litter sizehow many offspring should anorganism, say a bird, have in a particular litter? (11) Large littersmandatedecreased investment in individuals,among other costs,but increase the number of lottery tickets in the evolutionarysweepstakes. This problem has relevance across the taxonomicspectrum, and especially from the production of seed by plantsto the litter sizes of elephants and humans. Even for vertebrates,the evolutionary resolution shows great variation: The typicalhuman litter is a single individual, for which parental care is high, whereas fish may produce millions of offspring with low indi- vidual probabilities of survival.

    E C O N O M I C

    S C I E N C E S

    L U T I O N

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    51www.faculty.faireld.edu/faculty/hodgson

    Based on Survey of Consumer Finances

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    Extensions (with Ricky Der and others)

    Modify assumptions to try to produce Pareto tail Number of offspring contingent on wealth Non-uniform discounting among offspring Other sources of uncertainty

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    Extensions (with Ricky Der) Modify assumptions to try to produce Pareto tail Number of offspring contingent on wealth Non-uniform discounting among offspring Other sources of uncertainty Prosociality to non-relatives

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    Indeed, inter-generational equity isonly part of the problem

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    i94.photobucket.com/albums/l96/carlalynne

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    Also need to consider intra-generational equity

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    dericbownds.netSao Paolo

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    Inequity in the distribution of wealth

    is increasing

    56www.epi.org/page

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    Moreover, we live in a global commons,in which

    www.centerstage-musicals.com

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    Moreover, we live in a global commons, inwhich

    www.enn.com/news/enn-stories/2001

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    This is exaggerated when theindividual agents are nations

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    Environmental Kuznets Curve

    Shift industries in developed countries to lesspolluting ones

    Shift pollution and environmental damage todeveloping nations

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    http://www.marketobservation.com

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    The challenge.achievingcooperation at the global level

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    Prototypical problem:Prisoners dilemma

    Cooperate

    C o o p e r a t e

    Defect

    D e

    f e c t

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    Only stable solution: Nashequilibrium

    Columbia.eduCooperation loses

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    William Forster Lloyd (1832)

    Aelbert_Cuyp

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    But cooperation does arise inNatureand in theory

    peoplesgeography.les.wordpress.com

    How?

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    The evolution of altruism andcooperation

    morningnoonandnight.les.wordpress.com

    Delayed publication of Origin of Species for twenty years

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    Well understood:W.D.Hamilton and the social insects

    www.csiro.au

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    Well, not as well-understood as itused to be

    ANALYSIS

    The evolution of eusocialityMartin A. Nowak 1, Corina E. Tarnita 1 & Edward O. Wilson 2

    Eusociality, in which some individuals reduce their own lifetime reproductive potential to raise the offspring of others,underlies the most advanced forms of social organization and the ecologically dominant role of social insects and humans.For the past four decades kin selection theory, based on the concept of inclusive fitness, has been the major theoreticalattempt to explain the evolution of eusociality. Here we show the limitations of this approach. We argue that standardnatural selectiontheory in the context of precisemodels of populationstructurerepresentsa simplerand superior approach,allows the evaluation of multiple competing hypotheses, and provides an exact framework for interpreting empiricalobservations.

    For most of the past half century, much of sociobiologicaltheory has focused on the phenomenon called eusociality,where adult members aredividedinto reproductiveand (par-tially) non-reproductive castes and the latter care for the

    young. How can genetically prescribed selfless behaviour arise by natural selection, which is seemingly its antithesis? This problemhas vexed biologists since Darwin, who in The Origin of Species declared the paradoxin particular displayed by antsto be themost important challenge to his theory. The solution offered by themaster naturalist was to regard the sterile worker caste as a well-flavoured vegetable, and the queen as the plant that produced it.Thus, he said, the whole colony is the unit of selection.

    Modern students of collateral altruism have followed Darwin incontinuing to focus on ants, honeybees and other eusocial insects,becausethecoloniesofmost oftheir speciesaredividedunambiguously into different castes. Moreover, eusociality is not a marginal pheno-menon in thelivingworld. The biomass of ants alone composes morethan halfthatof allinsects andexceeds that of allterrestrialnonhuman

    greater thantwo timesthe cost tothe altruist( R 5 1/2) or eighttimesin the case of a first cousin ( R 5 1/8).

    Dueto itsoriginalityand seeming explanatorypower,kin selectioncameto bewidelyaccepted asa cornerstone ofsociobiological theory.Yet it was not the concept itself in its abstract form that first earnedfavour, but the consequence suggested by Hamilton that came tobe called the haplodiploid hypothesis. Haplodiploidy is the sex-determiningmechanism inwhichfertilizedeggsbecomefemales, andunfertilized eggs males. As a result, sisters are more closely related tooneanother( R 5 3/4) than daughters areto their mothers( R 5 1/2).Haplodiploidyhappens to be themethodof sexdeterminationin theHymenoptera, theorder of ants, beesand wasps. Therefore, coloniesof altruistic individuals might, due to kin selection, evolve more

    frequently in hymenopterans than in clades that have diplodiploidsex determination.

    Inthe 1960s and1970s,almostall thecladesknownto haveevolvedeusociality were in the Hymenoptera. Thus the haplodiploid hypo-thesis seemed to be supported, at least at first. The belief that haplo-

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    Relatedness is not the whole answer,or maybe not even the main answer

    Reciprocal altruism

    www.natureartists.com

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    Cooperation is easily explained in smallgroups, with repeated interactions

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    But how is cooperation sustained inlarger groups, like societies?

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    And can these principles be extendedto the global level?

    www.purdue.edu/envirosoft

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    Other contributors to prosociality Social norms (and enforcement)

    Societies, religions and governments Other institutions, like WTO or treaties

    eoearth.org

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    Social norms can sustain and enhanceprosocial behaviorE. Fehr

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    A theory for the evolution of other-regard integratingproximate and ultimate perspectivesErol Akay

    1,2, Jeremy Van Cleve

    1,3, Marcus W. Feldman, and Joan Roughgarden

    Department of Biology, 371 Serra Mall, Stanford University, Stanford, CA 94305

    Edited by Simon A. Levin, Princeton University, Princeton, NJ, and approved September 15, 2009 (received for review April 21, 2009)

    Although much previous work describes evolutionary mechanismsthat promote or stabilize different social behaviors, we still havelittle understanding of the factors that drive animal behavior prox-imately. Here we present a modeling approach to answer thisquestion. Our model rests on motivations to achieve objectives asthe proximate determinants of behavior. We develop a two-tieredframework by rst modeling the dynamics of a social interactionat the behavioral time scale and then nd the evolutionarily stableobjectives that result from the outcomes these dynamics produce.We use this framework to ask whether other-regarding motiva-

    tions, which result from a kind of nonselsh objective, can evolvewhen individuals are engaged in a social interaction that entails aconict between their material payoffs. We nd that, at the evo-lutionarily stable state, individuals can be other-regarding in thatthey are motivated to increase their partners payoff as well astheir own. In contrast to previous theories, we nd that such moti-vations can evolve because of their direct effect on tness and donotrequire kinselection or a specialgroupstructure.We alsoderivegeneralconditionsfor theevolutionarystability of other-regardingmotivations. Our conditions indicate that other-regarding moti-vations are more likely to evolve when social interactions andbehavioral objectives are both synergistic.

    Animalbehavior is determinedboth by proximatemechanismsthat dictate ananimalsactionsin real time andby evolution-

    ary forces that shape these proximate mechanisms. Even thoughtheevolutionarydynamics ofsocialbehaviorhavebeen extensivelystudied (14), proximate mechanisms of behavior and how theyinterface with evolutionary forces remain poorly understood (4).In recent years, some models have integrated a proximate mecha-nism with an evolutionary analysis (5, 6). Furthermore, an explic-itly two-tiered approach with potentially cooperative behavioraldynamics embedded in an evolutionary dynamic has been pro-posed (7)as necessaryto understandthe evolutionof social behav-ior.We contribute to thisliterature by developing a unied frame- work for modeling the evolution of a specic type of behavioralinteraction based on a well-dened proximate mechanism.

    Our proximate mechanism is based on the notion that ani-mals are motivated to achieve certain objectives. Goal-seeking

    behavior has been a recurring theme in animal behavior andhas been an integral part of earlier ethological thinking (e.g.8, 9). However, this idea lost its prominence after the emer-gence of modern behavioral ecology, which focuses mainly onthe tness consequences of behavior (see, for example, page 6

    payoffof itssocial partner, thefocal individual is said toexhibit another-regarding preference for its partner (12). We focus on suchother-regarding objectives for three reasons. First, the existenceof other-regarding preferences has received substantial supportrecently fromlaboratory experiments thatshow a capacityin somenonhuman primates for unsolicited food sharing even when therecipientcannotreciprocate(1315).Second,explanationsfor theevolutionof suchpreferencesare still in dispute (3,4, 16)and oftenrely on costly punishment and reproductive differences betweengroups (12) or on indirect selection on kin (13, 17) instead of on

    direct selection on the actions of focal individuals. Third, from aconceptual perspective, an other-regarding preference is a simple way in which the behavioral objectives of two interacting indi- viduals can be brought into concordance even when their payoff interests diverge. In this way, we can clearly delineate altruisticmotivations driving a specic behaviorfrom the underlying tnessconsequences of such behavior.

    Weintegrate the proximatemodel of behavioralobjectives withananalysisof theselectionpressures acting onthoseobjectives andnd two new results. First, we show that other-regarding objec-tives, and thus motivations, can evolve through direct selection onthe tness effects of individual behaviors. Second, we show thatsynergismin the payoffs fromthe social interaction and synergisminindividualsobjectivespromotethe evolution ofother-regardingobjectives. These synergisms are directly related to how the ben-ets and costs of different behaviors are turned into payoffs andhow individuals convert information about the payoffs from thesocial interaction into reward sensations.

    Results

    The Behavioral Model. We begin by developing a model of a socialinteraction in which two individuals share resources with eachother. Imagine, for example, two capuchin monkeys, one havingbeen given apples, the other carrots, as in the experiment by deWaal (18). Both individuals need the sugar in the apple and the -carotene in the carrot, so each would do best to exchange someof its holdings. For simplicity, we assume that costs and bene-ts of sharing are the same for both individuals. We label thedonation that a focal individual, individual 1(l1), makes to its

    partner, individual 2(l2), bya

    1 , and the donation that l2 makesto l1 by a 2 . We term a 1 and a 2 individuals actions and assumethat 0 a 1 , a 2 1. In the dynamical behavioral model below,the actions can be thought of as the rates at which individualsexchange donations. Suppose that the marginal benet a food

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    The Commons solution (Hardin)

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    The maintenance of cooperation in smallsocieties depends on shared and mutually

    agreed-upon norms

    Ostrom

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    In general, contributions to public goods/

    cpr depend upon

    Intrinsic prosociality

    Reciprocal arrangements and contracts Norms, laws, taxes and incentives

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    Examples

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    CPR/Public goods affect the value of

    private investments

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    y( x , Z ) = x Z Yield:http://www.travel-destination-pictures.com

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    Dixit and Levin:

    Prosociality and public goods

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    Dixit-Levin

    v 1 = u 1 + u ii = 2

    n

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    Dixit-Levin

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    Individual utility:

    where is prosociality , and < z > is the public pool,which benets from local prosociality and leakagefrom other groups

    v i = F ( x i , zi , < z > i ) + i k F ( x k , zk , < z > k )k i

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    Extension (with Dan Rubenstein)

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    These arrangements widespread in East

    Africa Maasai Samburu Turkana Boran

    mashriqq.com/?p=1081

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    Variety of mechanisms:

    Repeated game

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    Summary:

    Social/behavioral research topics Discounting and intergenerational equity Prosociality, public goods and common-pool

    resources Dynamics of social norms on networks Networks, contagion and robustness Role of leadership in behavioral shifts Institutions for achieving sustainability

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    Sustainability Research faces

    unique challenges Interdisciplinary and multi-disciplinary Human-environmental systems are complex

    adaptive systems

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    Ecological systems and socio-economic

    systems alike are complex adaptive systems

    90

    http://www.latinamericanstudies.org/maya

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    Adam Smiths Invisible Hand

    91

    www.bized.co.uk

    Th i i ibl h d d

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    The invisible hand does not protectsociety

    92

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    Those lessons are magnied for

    ecological and environmental systems

    93

    media-2.web.britannica.com

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    The CAS perspective means In both cases, management requires a balance

    between free-market and regulation

    New institutions must be adaptive Can adaptive features be built in? Robustness

    Trust and cooperation essential Key to macroscopic goals is in microscopic incentives Montreal Protocol?

    94

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    Can cooperation be extended to the

    global level?

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    Challenge is to integrate Bottom-up mechanisms, like evolved prosociality Top-down mechanisms, like rewards and

    punishments Collective actionTo achieve

    Adaptive, polycentric governance and agreements

    96

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    Need new institutions

    97

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    Need new institutions

    98

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    Climate change

    Polycentric approach (Ostrom)

    100pullingback.blogspot.com

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    Emergence of cooperation within

    groups is often for the benet of conict with other groups

    Lariviere

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    In the global commons, there is no

    other

    Understanding how to achieve international

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    Understanding how to achieve internationalcooperation is at the core of achieving

    sustainability in dealing with our commonenemy: environmental degradation

    103

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    Carole Levin