A stealth Templeton grant?

A new paper in Proc. Nat. Acad. Sci. USA by Alexander Stewart and Joshua Plotkin (link and citation below) has gotten some press (see, for instance, this summary in Archaeology News), for the results seems to show that evolutionary “game theory” will produce cooperative behavior.  People love that because it shows that selfish genes can produce unselfish behavior. (The Templeton Foundation also loves it because it creates a kind of fusion between evolution and niceness.)

Well, we knew about the evolution of cooperation already, but “game theory” analysis, in which different game-like strategies (usually involving two players) compete against other to see which strategy “wins” in a population, has a particular fascination for people.  Many of us know about the “Prisoner’s Dilemma” game, and the “tit-for-tat” strategy that can also produce initially cooperative behavior in such a game.

Game-theoretic studies of cooperation continue to appear, as people want to see why and how cooperation could have evolved in our species when it seems easier to imagine that selfish genes produce selfish behavior. I won’t try to refute the latter misconception here: Dawkins has done it many times before, and you should already know about reciprocal altruism, kin selection, and other situations in which genes producing cooperation have higher fitness.  And I’m dubious about such studies anyway, because they rarely have a genetic model behind them. If you want to show whether a complex behavior can theoretically evolve, you really need to have some genetic assumptions behind it.  Is a switch between two complex strategies due to a single gene? If not, how can you go from one complex behavior to another (which is how these models work) if many genes are involved, and the hybrid form may have an intermediate strategy.

But what I want to highlight here instead is the funding of the Stewart and Plotkin work. I won’t go into the gory details of their results, but will merely give the “layperson’s summary” of the paper:


Cooperative behavior seems at odds with the Darwinian principle of survival of the fittest, yet cooperation is abundant in nature. Scientists have used the Prisoner Dilemma game, in which players must choose to cooperate or defect, to study the emergence and stability of cooperation. Recent work has uncovered a remarkable class of extortion strategies that provide one player a disproportionate payoff when facing an unwitting opponent. Extortion strategies perform very well in head-to-head competitions, but they fare poorly in large, evolving populations. Rather we identify a closely related set of generous strategies, which cooperate with others and forgive defection, that replace extortionists and dominate in large populations. Our results help to explain the evolution of cooperation.

One of the authors gives a pretty strong statement about the significance of the work in the Archaeology News piece:

“Ever since Darwin,” Plotkin said, “biologists have been puzzled about why there is so much apparent cooperation, and even flat-out generosity and altruism, in nature. The literature on game theory has worked to explain why generosity arises. Our paper provides such an explanation for why we see so much generosity in front of us.”

And a blurb for the piece even appears on the Dawkins Foundation website.  But in truth I doubt whether this is a viable explanation for human generosity, which can have many sources besides a particular game-theoretic model.

What interests me more is who funded this work, and how the funding was mentioned.

If you go to the acknowledgments section of the paper, you will see a nod to funding from “The “Foundational Questions in Evolutionary Biology Fund”:

Screen shot 2013-10-06 at 5.29.19 PMWhat fund is that? I hadn’t heard of it before.  But if you Google it, you’ll be taken to a link on Harvard University’s server describing not a fund but a grant, and there you will discover that the bankroll for that grant comes from the deep pockets of—yes, you guessed it—The John Templeton Foundation. And if you click further, you’ll find that that grant is to the tune of $10,500,000, and the project leader is Martin Nowak, a religious evolutionist whom I’ve discussed before.

Now I’m not suggesting at all that the study’s results are tainted by Templeton’s funding. But what I am suggesting is two things. First, that Templeton gives out an inordinate amount of money for this kind of work, and that they know what kind of results they want. And if you produce those results, Templeton gives you more dosh, and you keep your seat on the gravy train. 10.5 million dollars is much more than an average National Science Foundation grant, and Templeton’s penchant for the numinous, and for the fusion of science and faith, deeply corrupts the research efforts of evolutionary biology by allowing researchers access to a huge pot of money with an agenda behind it.

Second, why did the authors not thank the Templeton Foundation directly? Could they be embarrassed or ashamed by the association? (I know I would be!) And had I been money-hungry enough to have taken that money to fund my work, I would have cited the funds honestly, as “A grant on the Foundational Questions in Evolutionary Biology from the John Templeton Foundation.” For there really isn’t a “Foundational Questions in Evolutionary Biology Fund.” There is a grant funded by Templeton. Templeton’s name doesn’t appear.

I don’t know if the investigators are ashamed of their funding, but I do wonder why they chose to disguise its source.


Stewart, A. J., and J. B. Plotkin. 2013. From extortion to generosity, evolution in the Iterated Prisoner’s Dilemma. Proceedings of the National Academy of Sciences 110:15348-15353

h/t: Todd


  1. Richard Olson
    Posted October 12, 2013 at 10:08 am | Permalink


  2. gbjames
    Posted October 12, 2013 at 10:19 am | Permalink


  3. Diana MacPherson
    Posted October 12, 2013 at 10:21 am | Permalink


    • Steven Obrebski
      Posted October 12, 2013 at 10:26 am | Permalink

      What does the comment “sub” mean??

      • Posted October 12, 2013 at 10:39 am | Permalink

        It means that people are subscribing, so that they can be notified of new comments. There is no way to be notified without posting a comment oneself.

  4. Posted October 12, 2013 at 10:44 am | Permalink

    Lovely comment on the funding. But sad.

  5. Posted October 12, 2013 at 11:11 am | Permalink

    Templeton’s reputation is preceding it.

  6. Posted October 12, 2013 at 11:20 am | Permalink

    What I find puzzling about these type of studies is that the authors assume that the behavior or the gene for the behavior is passed on to offspring without the slightest mention of Mendelian patterns of inheritance or gene theory. The authors assume “cooperation” is a trait that is handed down carte blanche. If you realize that these traits are expressed as dominant or recessive alleles then you can have the existence of maldaptive recessive traits or incompletely dominant traits like cooperation and explain their persistence in the same way scientists have explained the persistence of sickle cell alleles. In other words it might be bad to have two copies of a “cooperation” gene, but be good to have only one copy. The allele would be exploiting itself because the heterozygotes would benefit by having cooperative people in their midst, but would not be (or only be slightly) cooperative themselves and thus not be at a disadantage. Food for thought from an amateur geneticist.

    • BillyJoe
      Posted October 12, 2013 at 8:24 pm | Permalink

      I’m not sure that single gene determination of cooperative behaviour has much legs.

  7. Posted October 12, 2013 at 11:42 am | Permalink

    I admit that I am not that informed about all aspects of evolution, but it seems to me that the term altruism generally applies to humans and primates, and perhaps some other mammals. In any case one animal helping another animal without any immediate benifit to itself seems totally evolutionary, like a goose keeping guard while his flock feeds.

    • John Scanlon, FCD
      Posted October 13, 2013 at 5:37 am | Permalink

      I see what you did there, mr gosling.

      But any definition of altruism (or whatever) that can be applied consistently to primates will be found to apply much more broadly, without implying or assuming primate-like or human-like experience… or the existence of souls with free will.

  8. Posted October 12, 2013 at 11:59 am | Permalink

    I don’t see any value in becoming critical about the value of Evolutionary Game Theory in theoretical biology just because the Templeton Foundation has jumped on the bandwagon and is trying to use the subject as a touchpoint for it’s own particular pro-religious agenda. It is truly sad that Martin Nowak, who is one of the most brilliant mathematical minds in the subject area, is also a highly committed Christian, and Nowak seems exceptionally happy to swallow Templeton millions to say some nice things about religion. It’s all pretty shabby and disgusting, but it doesn’t throw out of the window the basic usefulness of the subject as a model for Evolution. There are numerous examples of behavioural studies that show that evolved animal strategies match EXACTLY the predictions derived from Evolutionary Game Theory and totally demystify these behaviours. This includes cooperative behaviours. As for having a complete genetic mechanism presently identified for these heritable behaviours, we are not there as yet. But that doesn’t negate the fact that they can exist. We really must be very very careful not to trash good science just because Templeton is throwing money at some of it’s researchers.

    • Posted October 12, 2013 at 12:07 pm | Permalink

      Excuse me, but I made it explicit that I was not criticizing the research because it was funded by Templeton. Did you even read that post? My explicit criticisms of game theory rested on its lack of explicit genetic underpinning. But I said that what I wanted to talk about more particularly was the researchers’ apparent desire to hide the source of funding. So the post has two distinct topics, and, if you read it, you would have seen this sentence:

      Now I’m not suggesting at all that the study’s results are tainted by Templeton’s funding.

      So your first and last sentences mischaracteerize what I said, and if you’re trying to admonish me, I expect you’ll take it back.

      • Posted October 12, 2013 at 1:00 pm | Permalink

        No admonishments intended.
        But your throwing cold water on Evolutionary Game Theory because we presently lack a specific genetic explanation of how these behaviours are coded DOES have the effect of negating this area in the eyes of the regular reader, especially given your stature in the subject area. Take for instance Torbjörn’s comment posted below:
        “ Thank you! I’ve been unsettled by use of game theory, and now I see why: it is a (sometimes) easily solved toy model, but not decisive.”
        Well, it’s NOT a toy model – it’s among the best mathematical work in the entire field of theoretical biology – and it is extensively supported by experimental results in measured behaviours, so it’s not just theory.
        And so what I’m trying to say, is that even if there is a gap in having the genetic mechanisms specifically identified doesn’t mean Evolutionary Game Theory isn’t a proper explanation. It just means there’s a gap still left to fill.

        • Diana MacPherson
          Posted October 12, 2013 at 1:43 pm | Permalink

          I hope you are not implying that readers here are incapable of nuance. I didn’t see Jerry’s comments as “throwing cold water on Evolutionary Game Theory” but instead relating why some studies should be questioned as proving an evolutionary reason for such behaviour. He wasn’t dissing game theory altogether and I thought this insight was helpful.

          • Posted October 12, 2013 at 3:38 pm | Permalink

            Honestly Diana, I really can’t see that there can be any questions whatsoever about the applicability of Evolutionary Game Theory (EGT) as the basis for explaining evolved behaviours. First, we must admit that we do observe in all species that behaviours ARE inherited, are innate …. that “like begets like”. The only mechanism available for this heritability MUST have a genetic basis. These behavioural “strategies” MUST be evolved, for there is no alternative derivation (unless you are a creationist). Now which competing optional (mutant) strategy will be most fit, will have the best chance of survival? Evolutionary Game Theory shows exactly what that strategy will be for any given level of cognition in particular circumstances (games). And nature mirrors this – what we find in nature is what EGT predicts. Yet we don’t know the exact genetic mechanisms for encoding this behaviour. Does this really call EGT into question? No, it is research to be done.
            Anyhow, back to the main point of the posting – the corrosive effects of Templeton money on the direction and interpretation of scientific research. On this matter, I think, we are all in total agreement.

          • Posted October 14, 2013 at 5:21 am | Permalink

            The difficulty is that the judgement of an expert has been presented, with no backup reason for further research. The utility of the results has been discredited without explanation, and that is not fair. Especially when it seems to these two gentle readers that game theory is useful in understanding why altruistic traits became widespread in the first place!

            All we want is an explanation, or else a slightly less one sided presentation (e.g. I’m not sure about their results, but I’m interested not in…). We respect your opinion, so if we are wrong we’d at least like a link to something explaining why, because we know that generally the opinions expressed here are well judged.

  9. Torbjörn Larsson, OM
    Posted October 12, 2013 at 12:22 pm | Permalink

    Oh dosh – they get ($10 MUSD) more creationist dosh for inanities of inserting religion into science than Russell got ($8 MUSD) from NASA to explore abiogenesis.

    [Which is a real Foundational Question in Evolutionary Biology!

    Not for the theory of course [shush, creationists, go home and play with soft toys instead!], but for the conditions of the chemical evolution process and the (initial) conditions of the biological evolution process both.]

    And I’m dubious about such studies anyway, because they rarely have a genetic model behind them. If you want to show whether a complex behavior can theoretically evolve, you really need to have some genetic assumptions behind it. Is a switch between two complex strategies due to a single gene? If not, how can you go from one complex behavior to another (which is how these models work) if many genes are involved, and the hybrid form may have an intermediate strategy.

    Thank you! I’ve been unsettled by use of game theory, and now I see why: it is a (sometimes) easily solved toy model, but not decisive. No wonder applied mathematicians have taken to it!

    • Latverian Diplomat
      Posted October 12, 2013 at 12:40 pm | Permalink

      What a bizarre thing to say in that last paragraph. Not only is applied mathematics one of the most difficult of intellectual pursuits, but one of the architects of the neo-Darwinian synthesis was Ronald A Fisher, who could best be described as working in the application of mathematics to the understanding of population genetics and evolutionary biology. (God knows the biologists in Fisher’s day needed the help with what can be some very hairy maths).

  10. Diane G.
    Posted October 12, 2013 at 12:24 pm | Permalink


  11. Latverian Diplomat
    Posted October 12, 2013 at 12:34 pm | Permalink

    Certainly this work can be oversold. But if nothing else it refutes the naive intuition stemming from the original Prisoner’s dilemma work that selfishness tends to be broadly optimal. Similar work has has also predicted conditions where a set of multiple strategies within a population is more stable than any single strategy for 100% of the population, which is an outcome that I believe has been observed in the wild.

  12. Posted October 12, 2013 at 3:44 pm | Permalink

    Also, for some context, there is this:http://www.ncbi.nlm.nih.gov/m/pubmed/?term=Nowak+MA%5BAuth%5D+AND+Plotkin+J%5BAuth%5D

  13. Posted October 12, 2013 at 4:25 pm | Permalink

    Thank you for taking the time to investigate the fine print of the ‘Acknowledgments.’ God works in mysterious ways.

  14. Posted October 12, 2013 at 6:15 pm | Permalink

    I am not sure why any one would try to attribute this to “shame” at all: this page shows clearly that the FQEB program invites grant proposals from other researchers, and given that the authors provided a grant number, it seems clear that they were referring to a grant obtained through this channel.

    I don’t agree with all that the John Templeton Foundation does, but admonishing the authors for this point seems to me to be not very different from admonishing someone for neglecting to mention that their NSF grants were funded by the US Government.

    • steve oberski
      Posted October 12, 2013 at 7:17 pm | Permalink

      It think that there is a direct parallel between what Templeton is doing with their “Foundational Questions in Evolutionary Biology Fund” and what the pharmaceutical industry does when it funds ghostwritten academic papers.

      In both cases the intention is to hide the connection between the author of the paper and the source of funding.

      For an interesting looking into how this plays out in the pharmaceutical industry I recommend
      “Bad Pharma: How Drug Companies Mislead Doctors and Harm Patients” by Ben Goldacre.

      • Posted October 12, 2013 at 9:41 pm | Permalink

        I haven’t read the book, so I apologize in advance if my reading of its Wikipedia summary is grossly inaccurate. With that out of the way, the allegations in Goldcare’s books seem much more sinister: that of papers bewing ghostwritten, presumably while completely hiding the original authors. On the other hand, in this case, the authors clearly mentioned the primary source of their funding in the usual place (and one simple search reveals the secondary source as well), so I don’t think the two situations are quite comparable.

  15. Posted October 13, 2013 at 4:40 am | Permalink

    I took the opportunity to have a look at the FQEB website (http://www.fas.harvard.edu/~fqeb/) and I’ve got to say that nothing has ever triggered such a combination of simultaneous enthusiasm and revulsion in me before.
    -Enthusiasm in the fact that there is substantial funding now available for scientific research into some really fascinating central topics in evolutionary biology that ought to be addressed.
    -Revulsion with the fact that combined with these topic areas there are other “research” sponsorships delving into areas of pseudoscientific religiously slanted claptrap. This produces the worry that some people will not recognise the difference between the good science and the claptrap or that the good research will be tainted by the source of sponsorship.
    Equally sickening is the way Nowak is introduced on the site home page as one of “The 50 Smartest People of Faith”. All I can say is, if these are the 50 best names that can be listed, I really wouldn’t bother featuring my own membership among them with such prominence.

    • Posted October 14, 2013 at 6:22 am | Permalink

      I think this should be compared to FQXi where the physicists have found a way to extract money from the Templeton Foundation without being compromised and pandering to religion.

      It was basically achieved by the physicists insisting that atheist physicists such as Anthony Aquirre in its Directorate and Advisory panels. This is why prominent atheist physicists such as Frank Wilcek, Sean Carroll, Lenny Susskind et al participate in it.

      It’s most recent discussion that verges on religion is a video by Sean Carroll on why God is a bad idea.


      • Posted October 14, 2013 at 11:18 am | Permalink

        Hmmmm. I just don’t know…. I can’t help having the impression that taking any Templeton money is entering a sort of whoredom. And yet…

        On the other hand looking at FQXi, which I hadn’t heard about till you mentioned it, I cannot fault the value of having funding for any of the research projects sponsored in their recent grants. The same goes for MOST of the topics from FQEB. I suppose one could argue from purely a cynical political position that it’s far better to take Templeton funds and use them for REAL science, than refuse to touch it and see it used only for the claptrap projects that prop up religion and do nothing to further scientific knowledge. But surely we can’t be naïve about how Templeton will exploit our involvement to meet their own agenda. Only someone with a good grasp of science can make a nuanced judgement about the scientific validity of a Templeton research project – the public at large can’t do this differentiation, so it’s Templeton that wins this little political game – the compatibility of science and religious belief is validated. It’s a damn high price to pay for furthering scientific knowledge.

  16. prasad
    Posted October 14, 2013 at 10:20 am | Permalink

    “But what I am suggesting is two things. First, that Templeton gives out an inordinate amount of money for this kind of work, and that they know what kind of results they want.”

    For once I’m not seeing a nefarious purpose when it comes to Templeton. This kind of research into the mathematics and evolution of cooperation only undercuts the notion that you need God for morality. That Templeton would fund it redounds to their credit.

    • prasad
      Posted October 14, 2013 at 10:20 am | Permalink

      Also sub.

  17. Joshua Plotkin
    Posted November 19, 2013 at 8:43 pm | Permalink

    “Why did the authors not thank the Templeton Foundation directly?”

    In answer to your question, the statement in the Acknowledgements section of our paper is a verbatim copy of the required statement given in the Grant Agreement. In other words, we acknowledged the grant precisely how the grantor required that we acknowledge them.

    Do you feel that any statement in the research paper is somehow a mis-representation of the work? In the “Significance Section”, meant for a general readership, we say simply that “Our results help to explain the evolution of cooperation.” I certainly agree that human generosity can have many sources or explanations besides a particular game-theoretic model; and don’t think our paper says otherwise.

    • gbjames
      Posted November 20, 2013 at 6:04 am | Permalink

      That is interesting. In other words the stealth was built in by Templeton’s desire to remain behind a curtain?

      Having complied with the required wording in the Acknowledgments section, was there anything preventing you from ALSO stating that the “Foundational Questions in Evolutionary Biology Fund” is an arm of the Templeton Foundation?

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