What evolutionary biology looks like to an economist

Just stumbled across this old (1996) speech of Nobel Laureate economist Paul Krugman’s, comparing the fields of economics and evolutionary biology. I always find it interesting and valuable to see how outsiders see a field I know as an insider. I think that Krugman paints an impressively-recognizable picture of evolutionary biology as evolutionary biologists know it, although I think these days there’s probably more emphasis on nonequilibrium dynamics in evolutionary biology (and ecology) than there was back in 1996. Very interesting read.

31 thoughts on “What evolutionary biology looks like to an economist

  1. Hah – a great read.

    Several impressions.
    First Krugman really is one of those rare truly bright guys who can understand a whole other field better than many who are in it.

    Second – what a great send up of Gould (and Galbraith in parallel). Also a great line about both fields requiring being good at algebra but full of people who hate algebra.

    Third – I love how he gets that all the interesting and truly profound evolutionary theory does center on natural selection leading to appropriate optimization (with constraints, possible frequency dependence, etc). People like Hamilton indeed don’t worry about mutation rate limitation, genetic load, neutral molecular evolution, hitchhiking, etc. But if you read the journal Evolution you would think these were the most important. I wonder if Krugman knew that and conveniently ignored it to take to task evolutionary ecologists or if that is something that only comes out if you read the primary literature (and follow “the algebra” as Krugman called it). I think he knows because he acknowledges two cases in economics where the journey is more important than the endpoint (and I acknowledge these exist in evolution too).

    Fourth – I find it interesting economists are so open about saying I come from the Marxian camp or the neo-Keynesian or Galbraithian or etc camp. We certainly have such threads and camps in our thinking in ecology and evolution but we aren’t supposed to openly acknowledge them as they would perceived as blinders to objectivity. Of course modern philosophers of science (e.g. Lakatos and beyond) all know that the economic approach is closer to how we really work. We’re just fooling ourselves when we don’t admit to belonging to an intellectual camp.

    • Re: your first impression, yup.

      Re: your second impression, I’ve quoted Krugman’s snark about Gould in my old post shredding “Spandrels of San Marco”, but I didn’t know where it came from until I found this speech yesterday.

      Re: your third impression, I agree with that point but not everybody would. John Whitfield once did a series of blog posts about Darwin’s Origin, in which he noted that you could group major evolutionary theoreticians into two rough schools. The “lean and mean” school focus on natural selection, use deliberately simple, elegant models, and claim Darwin as their intellectual ancestor. They’re also mostly skinny British men. 😉 Think folks like Fisher, Hamilton, Maynard Smith, Price (who moved to Britain), with Dawkins as their popularizer. Marek Kohn’s A Reason for Everything is a very good book (part history, part potted biographies) about this school of thought. The opposing “let a thousand flowers bloom” school, exemplified by Gould, maybe Sewell Wright, and possibly (as you’ve argued) many “ordinary” evolutionary biologists, considers selection just one among several evolutionary forces and focuses on the complex interplay of those forces. Interestingly, this school also claims Darwin as it intellectual ancestor, and not without reason. I’m a “lean and mean” man myself, and I take it you are too. If you don’t have evolution by natural selection, you don’t have lots of living organisms to exhibit drift, migration, mutation, etc. So all that other stuff is secondary in a very important sense.

      Jeremy Yoder (evolutionary biologist postdoc who writes the Denim and Tweed blog and for Nothing in Biology Makes Sense) has an old post arguing that genetic drift is really underrated as an interesting evolutionary force. But I haven’t been able to find the post just now by casually googling…

      Re: your fourth impression, interesting, I never really thought of that. Of course, for us ecologists and evolutionary biologists to openly acknowledge our “camps”, we’d have to have names for them. The fact that we don’t is probably evidence for your point here. Though I wonder a little if the camps are well enough defined in ecology and evolution to merit names. Worth noting as well that different camps in economics are associated with different graduate programs. Krugman is a “saltwater” guy, meaning he got his PhD at one of the name programs in a coastal state (places like MIT and Princeton). His opponents are “freshwater” guys, who got their PhDs at Chicago and Minnesota, near the Great Lakes. Maybe one reason we don’t have named “camps” in ecology and evolution is because we don’t have graduate programs that quite self-consciously teach one and only one school of thought.

      • Freshwater ecologists do talk about “camps” to some extent, though I don’t think I’ve heard that particular term used. But there are people from the Hutchinsonian perspective, from the Birge/Juday/Wisconsin perspective, and, as I believe David Post argued at the Cornell Limnology Celebration a few years ago, there is also a Needham/Cornell that is sort of intermediate between Hutchinson and Birge/Juday.

    • I’m sincerely flattered by that suggestion. That is the first, and probably last, time the phrase “you and Krugman” has ever been used to refer to me. 😉

      Exactly what sort of evolution & economics workshop might be worth doing would need a lot more thought. Perhaps something inspired by the ongoing economics crisis (“What major perturbations and regime shifts teach us about ecological and evolutionary dynamics”)? And honestly, I don’t think I’d be the best person to organize such a thing.

  2. “I have even tried to talk to some of the biologists, which in this age of narrow specialization is a major effort.”

    Indeed. At times, I’ve even gone so far as to try to understand them. My bad!

    Anyway, good post.

  3. Any reading suggestions for an econ grad student who is interested in learning more about the non-equilibrium dynamics you mention?

    • Hmm, good question. There’s now a very large theoretical literature on what’s known as “adaptive dynamics”. Adaptive dynamics is a particular modeling approach related to, but not identical with, older approaches like ESS models and quantitative genetics models. Michael Doebeli just wrote a book (for the Princeton Monographs in Population Biology series) on his adaptive dynamics work. Peter Abrams is another big name in the adaptive dynamics literature, and has probably focused on nonequilibrium dynamics more than most other theoreticians in this area. I have a couple of papers with David Vasseur in American Naturalist using this approach to reveal interesting transient dynamics on the way to equilibrium. And Steve Ellner and Nelson Hairston Jr. and their collaborators have a really great body of modeling and experimental work on the coevolution of predators and prey and how that affects predator-prey cycles. I’m sure there’s other stuff out there, but I’m not actually a proper evolutionary biologist–calling myself an evolutionary biology “insider” is a bit of an overstatement, honestly! So there may well be other good stuff I’m not thinking of or am not aware of.

    • Shameless self promotion but actually on target – I have an Annual Review of Ecology Evolution and Systematics 2007 paper (Evolutionary Game Theory
      and Adaptive Dynamics of Continuous Traits) that is intended 100% as an introductory guide to evolutionary game theory/adaptive dynamics (snuck it in as a review but its really a tutorial) that I co-wrote with Joel Brown that will give the biologist’s perspective. Definitely not in the league of the sources Jeremy mentions but as a starting point it might be most accessible.

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  5. For what it’s worth, it seems to me that Krugman’s swipe at Gould and Eldridge says more about who he’d been talking to in the field than any objective appraisal of opinions about the state of evolutionary biology in 1996. The “evolution by jerks” comment is just setting up a straw man. A different author with a different mindset who had talked to Gould’s “camp” would no doubt have used the “evolution by creeps” quote. Ultimately it was two groups of evolutionary biologists, both of which made important contributions to our understanding of pattern and process, talking past each other because their focus was different: micro- versus macro-evolution, if you like. Krugman doesn’t seem to have understood this.

    Gould and Eldridge made no secret of the fact that they were fundamentally left leaning in their political views and this seems to have annoyed Krugman (and those biologists he was speaking to) more than any real appraisal of their contribution.

    • As is obvious from my comments, I don’t agree with you, but I don’t have much to add to what I’ve written. That there were different camps doesn’t mean they were merely different.

      In terms of where Krugman’s views come from, you’re incorrect if you think Krugman got his views just from talking to people only in one camp. I don’t know who all he’s talked to, but he’s also read widely, including reading much of Gould’s work. So Krugman’s views do not reflect him being unfamiliar with, or only getting a biased presentation of, the views of one camp. And with respect, you’re wrong in thinking Krugman’s views reflects politics. Krugman’s a leftie himself, although not a Marxist. And Maynard Smith, someone who Krugman would see as at the core of evolutionary biology, was a Marxist like Gould.

      You might be interested to check our Marek Kohn’s book A Reason for Everything, which includes potted biographies of many of the leading figures of Fisher, Haldane, Price, Hamilton, Maynard Smith, and others at the core of the selection-focused school of evolution, and makes clear that there is no correlation between one’s political or religious views, and one’s views on natural selection. Those folks all saw pretty much eye-to-eye on natural selection, but were massively different from one another in almost every other way.

      • I’m not sure how you conclude that Krugman was speaking to a wide range of people when he makes statements regarding Gould such as “informed readers eventually conclude that there’s no there there” or “Gould’s ideas….[giving]…..an almost completely misguided view of where the field is and even of what the issues are”. Clearly Krugman was not talking to the many scientists who cited Gould’s work in a positive way. Spandrels, for all its shortcoming (and let’s not forget that it was written over 30 years ago and hindsight is always perfect) has been cited 2369 times to date. Not all of those referrals are pointing out its failings. Krugman’s just plain wrong when he implies that “real” evolutionary biologists thought that Gould talked crap. The only way he could come to that conclusion would have been if he had spoken only to Gould’s detractors. And many of them seemed to be motivated by jealousy as much as anything.

        What seems to be missing here is any acknowledgment that Gould’s training and expertise was primarily in palaeontology. Immediately that’s going to give him a different perspective on the evolution of life’s diversity from someone with a background in population genetics. That doesn’t make his perspective wrong as Krugman seems to be saying. It just means it’s different. Krugman’s reading of the field seems to be skewed in that he interprets “evolutionary biology” only in a way that is analagous to economics and ignores those aspects of the (broad) field that don’t fit the story he’s trying to tell.

        I agree, the politics is probably a red herring, though Gould never described himself as a Marxist -his detractors did, and the mud seems to have stuck. And if Wikipedia is correct, Krugman sees himself as a Liberal, in the American sense, which can mean a lot of things. Thanks for the book tip off, I’ll take a look.

      • I think you’re mistaking Krugman’s typically-pointed language for him being biased or uninformed. Krugman is well aware of how widely Gould has been cited, etc. If you haven’t done so, you may wish to read his blog and NYT columns (he’s in my blogroll). You will see that his language is equally strong when criticizing economic and political opponents with whose work he has intimate, first-hand familiarity.

        Krugman and those who agree with him on his views of evolutionary biology (like Brian and I) could of course be wrong (it’s always possible that anybody is wrong about anything). But no, it’s not because we’re uninformed or only talking to the wrong people or jealous or driven by politics or whatever.

        I’m guessing you’re an admirer of Gould; if so, that is absolutely fair enough. I don’t at all share that admiration, but I certainly don’t assume that it comes from you being biased or uninformed or jealous or a leftie or etc. With respect, I think you’re much too quick to assume, based on someone disagreeing with you and saying so using strong language, to infer that they must be biased or uninformed or have bad motives.

      • p.s. The fact that an idea or person has been widely cited as correct does not make it or them correct, or even partially correct. The intermediate disturbance hypothesis has been widely cited, and it’s in all the textbooks. It also has little-to-no empirical support and is logically invalid.

      • It’s one thing to criticise using pointed language but it’s quite another to claim to have inside knowledge of how a whole field perceives an individual’s work, which is what Krugman was doing in that speech. He can’t possibly have been, ergo his opinion was biased.

        There seems to be no room for a nuanced, historical view of the contribution of Gould. It’s not about an idea or an individual being “correct” or “incorrect”, it’s about the contribution it/they make. Even if punctuated equilibrium, exaptations, the Wonderful Life hypotheses, and all the rest of it are complete nonsense then they have still made a contribution to the field by stimulating other individuals to generate the data and ideas to test them. That’s how science works. As you say, every scientist gets things wrong on occasion. That’s not to devalue their work as scientists.

        I have to say that I wasn’t suggesting that you personally were jealous of Gould’s success. But you must acknowledge that high profile scientists who achieve “popular” success ALWAYS come in for greater criticism compared to those who stay within the academy.

        As for your view of the IDH, I’m familiar with your arguments and look forward to reading the TREE paper.

      • We’ll have to agree to disagree on whether Krugman is biased, sorry.

        I freely admit that Gould stimulated others to do valuable science. But here’s the thing: if Gould hadn’t ever written a word, those others would still have done valuable science, just different valuable science. So I respectfully disagree that inspiring others to do good science somehow compensates for getting things wrong.

        To be clear, I’m not saying Gould was a bad scientist. We all do the best we can, and we all get things wrong. But when you get it wrong, you get it wrong, and your colleagues have not just a right but a professional obligation to say so. And when they say so they’re not “devaluing” your work, they’re doing their jobs. Further, if Gould gets criticized in strong language, well, he dished it out as well (see my old post on Spandrels). If you express yourself in strong language, you have to expect people to reply in kind.

        I don’t know if scientists who achieve popular success come in for more criticism from their less-publicly-famous colleagues or not. Yes, Gould has been widely criticized by less-famous academics. But what about someone like Jane Goodall? (that’s an honest question, I have no idea how Jane Goodall is regarded by less famous primatologists or animal behaviorists) As far as I know (which isn’t very far), anonymous academic physicists have huge respect and admiration for Stephen Hawking. I wouldn’t want to generalize from the very small number of examples with which I’m sufficiently familiar to judge.

  6. I don’t disagree with anything you say in that last post, Jeremy. You’re right, we as scientists have an obligation to critique our peers. But I also see a contradiction in your position. On the one hand you state that “’I’m not saying Gould was a bad scientist” but on the other you’re supporting Krugman who said exactly that in 1996, if not in so many words.

    Regarding scientists who achieve popular success, there’s some fairly vitriolic comments on the web, apparently from physicists. See for example the comments below this book review:


    Perhaps it’s mainly a British phenomenon? We seem to be good at belittling success.

    Likewise I have no idea regarding Goodall.

    Thanks for the interesting discussion; have a good weekend.

    • I don’t think my position is contradictory. I think Gould was very wrong about a lot of stuff. I think his rhetoric and style often covered up for shortcomings in his evidence and arguments (I think that’s what Krugman’s getting at when he says there’s “no there there”). I think his whole approach to science has serious limitations that aren’t as widely recognized as they should be (see my Spandrels post). But none of that makes you a bad scientist, I don’t think. It doesn’t indicate bias, or sloppiness, or ignorance, or incompetence, or dishonesty, or etc. Bad scientists are pretty rare, I think.

      The only respect in which I think Gould was a bad scientist was in fudging skull data in The Mismeasurement of Man so as to be able to accuse someone else of fudging those data. And in the absence of evidence to the contrary, I’m willing to assume that was an isolated thing.

      • I’m willing to go a bit further than Jeremy. The fairest obituary I saw of Gould was I think in the New York Times that pointed out that he generated many ideas that changed the direction of research and that nearly all of these ideas turned out to be wrong. I think those are both true statements, but I don’t think by themselves they innately make the person that description applies to a good or a bad scientist. I can think of other famous ecology and evolution people you can apply those words to accurately that I would actually call great scientists.

        My issues with Gould are on other fronts:
        1) He never admitted he was wrong even when there was a preponderance of evidence. Now a lot of scientists are not great at admitting they’re wrong, but most at least know when its time to start being quiet. Gould went the opposite direction and upped the rhetoric.
        2) I believe the ideas he generated were in fact strongly driven by his political beliefs – his entire career was devoted to tearing down the adaptationist paradigm. Whether this was done out of ego or political bias I can’t say for sure, but I strongly suspect political bias was part of it – he wasn’t the only Marxist-motivated scientist-to take on adaptationism (think Lysenko). But even if this wasn’t his motive his goal was wrong-headed. Who in the 2nd half of the 20th century could think the adaptationist paradigm was completely wrong and needed to be dismantled? And claiming he was just trying to clarify the edges and limits doesn’t hold water – he was way over the top for that. And he explicitly stated that his view of what he was doing was bigger than that.
        3) He was way too quick to go for rhetoric instead of produce data.
        To my mind these are fatal flaws in a good scientist.

        A personal anecdote (I took a large lecture course from him). He’s the only professor I know who would create a final exam with a question that says “is my theory revolutionary?”. (I argued for incremental improvement and was marked wrong – 0 credit).

      • I agree with 1 and 3. As I’ve said, I don’t think 2 holds up, at least not in a simple way. Again, remember that Maynard Smith was a Marxist. Haldane too. And Marx himself admired Darwin’s work. So whatever Gould’s motives might have been, I don’t think they can be summarized as simply as “Marxists hate adaptationism”.

        If forced to summarize my own view, I wouldn’t say Gould was a bad scientist. I’d say he was unique, and deeply flawed.

        That anecdote about Gould’s teaching is rather jaw-dropping. In all honesty, in all my years as a student and prof, I’ve never even heard of such a thing. I mean, I’ve encountered lots of bad teachers, and lots of arrogant ones. Heck, I have more than my share of arrogance, and I’m also far from the best teacher I know! But it takes a pretty special combination of badness and arrogance to set that sort of essay question (which any decent teacher would see as having no single definitive right answer) and then give zeroes–not just some marks off, but zeroes!–to students whose answers you disagreed with. I’ve heard other nth-hand anecdotes about Gould’s teaching that indicate that it was bad in other ways, but nothing like that. Wow.

  7. Brian – I have to concur with Jeremy on the political aspect, I think it’s a red herring. My original comment regarding Krugman’s bias was with respect to his perception of Gould’s politics, not on what Gould’s beliefs may have been.

    Regarding the idea that “nearly all of these ideas turned out to be wrong”, I think it’s far too early to make such a statement. The debates about frequency of punctuated equilibrium and phylogenetic affinities of the Cambrian Explosion taxa still continue amongst palaeontologists, for instance. Obituaries are the worst kind of peer review: there’s no possibility of the author writing a response 🙂

    The anecdote is amazing, I agree, and very revealing. There’s no doubt that Gould was an arrogant man, full of his own self importance. But it must be hard to keep your feet on the ground when you’re being told by the world how brilliant you are. It happens in show business constantly. I saw Gould lecture twice. The first time was in Oxford in about 1990 when he was promoting the Wonderful Life book and he was brilliant, a captivating speaker. I was just beginning my PhD research and I was in awe of him, I’ll admit. The second time was in about 1996 at a Royal Society Discussion Meeting in London. He gave an awful talk, reading direct from his manuscript and coming across as incredibly pompous. Something seemed to have changed in that intervening period, though an n=2 sample size is rather weak support for such an assertion!

    • “must be hard to keep your feet on the ground when you’re being told by the world how brilliant you are.”

      Maybe. But certainly, there are famous people do it (and conversely every prof can also name arrogant colleagues who aren’t famous at all). Gould’s hero Charles Darwin was hugely famous, but by all accounts a really nice guy.

    • Jeff & Jeremy,

      I agree that not all Marxist scientists drive their science from their politics (Russia got pretty far in the sciences …). But I haven’t heard any arguments why this was’t true for Gould as some individuals clearly did do this (I gave the example of Lysenko). Gould was very clearly driven by some agenda. I of course can’t marshal any really strong arguments for my belief that it was Marxism – its just intuition from listening to him lecture for 45 hours or so, but I don’t have a smoking gun.

      Regardless of motivation (I acknowledged earlier ego could be an alternative) I think setting out with an agenda of destroying adaptationism is a pretty clear form of bad-science to: a) be so agenda driven rather than results driven, and b) pick an agenda like that.

      Jeff your n=2 story is reminiscent. My college roommate took Gould’s course as a freshman and came back after every lecture raving about how brilliant he was and how he tied baseball to evolution and etc. I took his course in my last semester of my senior year (and sat next to a friend in the same stage) and we both found him so pompous that he was actually a bad lecturer – he spent more time being grandiose than communicating. I’m pretty sure my roommate would have thought the same thing if he took the course in his last year (this was 1984 vs 1988). Maybe Gould changed, but maybe one’s perspective changes too as one becomes more trained as a scientist. The BS detectors tend to become stronger.

      • You obviously have a larger and different information base on which to judge Gould than I do. Though I would suggest that it ought to be possible to judge the science and the motives independently. I don’t have an agenda to destroy the IDH–but if I did, I’d still be right about it. Although then again, I agree that the line between substance and motivation is fuzzy with someone like Gould, who relied so much on rhetoric to sell his ideas.

        Jeremy Yoder once said to me that he grew out of Gould–loved his writing in high school, but started seeing it as BS in grad school. I had much the same experience, and it’s my impression that that’s a pretty common experience among professional evolutionary biologists.

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