Beware of Stephen J. Gould

He wrote as if the entire Williams revolution had never occurred! Gould attacked, as if they were still current views, romantic notions that no serious biologist had put forth since the 1960s.

he pulled the same trick with mismeasure of man.

But some readers may have to take my word for this, since the names of eminent scientists are often less well-known to the general public than the names of fast-talking scoundrels such as Uri Geller or Stephen J. Gould.

actually, two things: 1) do a literature citation search.

2) appeal to another authority of some note, e.g., paul krugman saying the exact same thing re: gould & j.m. smith.

My impression is that Steven Wolfram got away with publicly claiming credit for a lot of people's ideas in his recent book "A New Kind of Science", in the sense that the public was never set straight. Gould probably could have too. My guess is that he simply didn't understand the logic and the conceptual background behind the "Williams Revolution", and was able to ignore it because it was unknown even to most professional biologists and to paleontologists like himself. Evolutionary biologists knew about it, but there aren't very many of them. From outside, if you don't understand their arguments, they can be ignored as one minor sub-specialty with some heterodox ideas obscured behind pretentious math. He didn't need neutronium earplugs. He just ignored his tiny band of critics with the same sort of justification, superficially, as I use when ignoring various specialties in macroeconomics. They are too small, demand too much thought to follow their arguments, and can't, as far as I can tell with minimal effort, make useful predictions. Probably, his idea of minimal effort is was than mine is, but after all, he was just a paleontologist who gained a lot of status. Status pretty inevitably reduces people's effective intelligence.

Gould's problem was basically that he didn't realize that evolutionary biologists were not some small sub-field, no more relevant than their size within biology and the historical sciences would suggest, but were rather the only scientists with expertise relevant to the questions he was discussing.

Thus we return to this blog's regular program, that of trying to figure out good general high-level heuristics for identifying the relevant experts on an given question.

Vassar, I think you're being far too charitable. Gould's critics were hardly a tiny band on any of the subjects he mangled. Once is accident, twice is coincidence, three times is enemy action, four times is stupidity, twenty times is either fraud or its moral equivalent in self-interested blindness.

But remember, evolutionary claims of progress were (and are) rife in the social sciences. That was what Gould ended up changing, if nothing else.

Surely we have many concepts of "complexity", not all of which directly translate into the number of fundamental bits you are discussing. So isn't there still room to argue whether these other forms of complexity are increasing at all, and if so whether via movement of mean complexity or of variance?

Robin: There are, but George Williams made his case for more than one form of complexity having not "progressed" over the last hundred million years, including complexity of morphological form, etc. So I stand by my statement that Gould utterly misrepresented what the field of evolutionary biology thought about complexity, creating false chaos so his bizarre criticisms could appear as avenging heroism.

The only progress I would defend as being visible in some recent evolutions is progress in quality of vertebrate brain software (not complexity or size per se), and this shift in adaptive emphasis must necessarily have come at the expense of lost complexity elsewhere. Look at humans: we've got no muscles, no fangs, practically no sense of smell, and we've lost the biochemistry for producing many of the micronutrients we need. This is yet another shift of adaptive emphasis, which in this case just happened to win the lottery, not an illustration of a general trend toward evolutionary progress.

There were eras of genuine evolutionary progress in some organisms, just not recently. They would occur subsequent to the development of new error-correcting mechanisms that shifted upward the equilibrium of adaptive complexity.

"progress in quality of vertebrate brain software (not complexity or size per se), and this shift in adaptive emphasis must necessarily have come at the expense of lost complexity elsewhere. Look at humans: we've got no muscles, no fangs, practically no sense of smell, and we've lost the biochemistry for producing many of the micronutrients we need." This looks suspicious to me. What measure of complexity of the brain's organization wouldn't show a big increase between invertebrates and humans? For the lost complexity you claim, only the loss of smell looks like it might come close to offsetting the increase in brain complexity; I doubt either of us has a good enough way of comparing the changes in complexity to tell much by looking at these features. If higher quality brains have been becoming more complex due to a better ability to use information available in the environment to create a more complex organization, there's no obvious reason to expect any major barriers to an increase in overall complexity.

Peter, I specified "within vertebrates" because many invertebrates - for example, bacteria - have lower-fidelity DNA copying, hence they may legitimately be expected to have a lower equilibrium level of sustainable complexity.

Eliezer,

You are in way over your head on this one. It is clear that you are behind on current literature. The "selfish gene" is old hat and out of date. Multi-level evolution is increasingly widely accepted by many geneticists, with such mechanisms as reciprocal altruism being keys. The equations for when the Williams argument on this matter breaks down have been around since James F. Crow first put them out back in 1953, although they were re-codfied and more widely distributed in the 1970s as the Hamilton-Price equations.

I am in my office where I do not have copies of the relevant books, and I never read the one by Gould that has you most worked up. It may well be that Gould claims excessive credit for ideas that were due to Williams in it. I do not know. I do know that while they disagreed on various things, Gould certainly does cite Williams in other places and writings and clearly recognized his role in various matters, at least in other places, if not in this particular book. There is little doubt that you are wildly exaggerating his sins in this particular matter.

Also, while one can find the idea of a variable rate of evolution in Darwin, there is no doubt that he saw evolution as fundamentally continuous and extremely gradual. "Nature non facit saltum" is the frontispiece for all editions of his Origin of Species, which comes ultimately from Leibniz. The guy you linked to dismisses Gould's (and Eldredge's) idea of "punctuated equilibrium" as already known to Darwin and thus derivative. Excuse me, but this is horse manure. Gould and Eldredge may have overstated the case, but they completely altered the discourse, and the awareness of the possibility of much more rapid evolution is now much more widely accepted.

Indeed, these ideas are linked, as I suspect you know. The possibility of more rapid evolution is indeed tied up with the idea of multi-level evolution, if only partially. It is indeed a "hardline, fundamentalist Darwinian" position that denies this, as it has been by Dawkins (and Williams earlier).

Barkley, since you haven't specified any of the reasons you think Williams's argument breaks down, I can't respond to that. The Hamilton/Price Equation deals with covariance, not mathematical information, and what does relatedness have to do with any of this? You would seem to be responding to something other than what I wrote.

As for multi-level selection invalidating the gene's eye view, it most certainly does not. It simply means that there are potentially vehicles of selection besides individuals. It doesn't change the distinction between vehicles and genes.

So I'll simply note that, while I am not a professional evolutionary biologist, it's not just me who thinks Gould is systematically misrepresenting evolutionary science. It's Richard Dawkins, John Tooby, John Maynard Smith, and Tooby's whole laundry list. I really don't think you can accuse that whole laundry list of failing to be up on the professional literature.

I don't object to the evolution content by any means - indeed, I find it interesting - but isn't attacking the work of one particular evolutionary biologist a bit off-topical to overcoming bias? (I had the same thought about the recent "congratulations to Paris Hilton" post - while it could have been used to point out the biases involved in signing/not signing up to cryonics, it didn't attempt to do anything of the kind.) Of course, you could argue that it's important to dismiss false information going around, in order to prevent people from being biased when thinking about evolutionary psychology... but to do so seems exceedingly narrow - if you want to debunk every person spreading false information about science, you won't have the time to write about abtual biases.

Eliezer, your non-response causes me to conclude that you aren't thinking clearly. John Maynard Smith's comments on Gould are adequate. Listen to Kaj and stick to areas where you know enough to be useful.

Gould's rhetorical strategy was straight from the Marxist book he was influenced by as a child: the status quo is reactionary, bad, and monolithic; revolutionaries are good; everyone is a vector of the Hegelian dialectic except 'us'. So he had no problem setting up caricatures as opponents, a bunch of speciecentric believers in progress and eugenics, biased beyond belief.

His Mismeasure of Man is a great example of his methods. His criticism mainly centers on pre-1940 work as if their sloppy methods and biases render human biodiversity disproven: By that logic, Pasteur, Mendel, Millikan, Flemming, Freud, Eddington all should have negative signs attached to anything they asserted (see Einstein's Luck). He employed the rhetoric of emphatic reassertion "Say it five times before breakfast tomorrow: … Human equality is a contingent fact of history". He dwelt on anecdotes, such as a question from a 1917 IQ test (Who was Christie Mathewson?), though there was subsequently a huge amount of research on much better tests when he wrote this in 1980. IQ is important, and you can measure it.

Straw men. Repetition. Ad hominem attacks. Totally ignoring the better arguments from your opponents. That was his style.

Eliezer,

Well, the statement from Tooby that you quote says exactly nothing precise. He refers to a "tangle," but nothing specific. Then he gives his shopping list of evolutionary biologists whom he considers more important/worthy than Gould. Fine, some of those certainly are and some others may be. This is hardly some slam dunk comment, but reeks all too much of the jealousy of the less popular for the more popular. I might as well get all bent out because Tyler Cowen and Bryan Caplan have written bigger selling books than I have.

Regarding Williams' argument, the issue is indeed the locus of evolution. Saying that it is the gene that gets passed on does not cut it. The question is the locus of activity that determines which genes get passed on. In the case of Dawkins and Williams this locus was generally (pretty much always in the case of Dawkins) the individual. Williams actually allowed for higher level selection as an occasional possibility. However, he ruled it out in general, and did so on essentially prisoner's dilemma, game theoretic grounds, the basis of Maynard Smith's position as well, the developer of the concept of the evolutionarily stable strategy. Maynard Smith was also very impressed with how easy it is for individuals to undermine cooperative strategies.

So, now we know better. Groups that cooperate can outcompete and replace groups that degenerate into internal back-stabbing and cheating. The evolution of such cooperation remains one of the really big questions out there. It turns out that the Crow-Hamilton-Price equations in fact give conditions for when such higher level evolutionary processes can become the dominating factor in determining which genes get passed on.

Regarding Gould, again, I am not going to defend him against all criticisms. I am not surprised that he has not always properly cited others. Certainly he had a large ego and did a lot of self-advertising. And I am perfectly willing to accept that he may have engaged in some oversimplifications or even obfuscations and confusions regarding evolution in terms of public knowledge. This is unfortunate to the extent it is true. But his debates with Dawkins and Williams and Maynard Smith were serious debates, regarding which on at least some issues the consensus is moving more in his direction now.

As for people dumping on his politicis, fine. But please do not pretend that you are making scientific arguments. These are essentially ad hominem arguments of the worst sort.

Barkley, read the rest of Tooby's criticism, not just the quote I excerpted for the particular purpose of showing the list of scientists. The link razib posted to Krugman is also directly relevant: You would seem to be one of the economists whom Gould managed to bluff into the impression that he was a major evolutionary theorist.

It's always nice to see OB pick out a specific and indubitable item of bias, adjust for range and wind, click the mouse, and convert it into a pink spray of little bias giblets. A really nice post. More of the same, please.

However, the question this post leaves unanswered is: given that Gould's reputation was mud in his own field, how come we've all heard of him? Why did I grow up on The Panda's Thumb and Mismeasure of Man?

Let me put it more directly: who do you have to know if you want to be Stephen Jay Gould?

After all, Gould was a good writer, but his eloquence was hardly unique among his competitors. For everyone who achieves the fame of a Gould, surely there are ten or fifteen people who could but don't. When you realize that Gould was a quack, the question's urgency rather increases. Who makes this decision? Is it just random? Or is there some predictable pattern?

If the latter, has this pattern affected any field except for evolutionary biology? How does it relate, if at all, to Lysenkoism? Can the two be considered homologous, or are they merely analogous?

Mencius: would you ask the same about Rowling? I think it's probably just an ordinary information cascade? To all those who have, more will be given.

Eliezer,

Well, let me see. Where shall I start?

Tooby is not an evolutionary biologist or population geneticist. He is an evolutionary psychologist. This screed from 1997 is about half a whine about Gould's treatment of some of his work. Perhaps the whine is justified, but it is a whine, and most of the rest of it is off-base. The argument about inverted panadaptationism is simply wrong, as a perusal of Gould's 1400 page plus magnum opus, The Structure of Evolutionary Theory, will tell. BTW, Tooby does not make it into the 43 page bibliography of that book, although Williams is discussed at great length and with great respect in it. I would note also that Williams altered his views somewhat between 1966 and his 1992 book. (Just for the record, I like the main book by Cosmides and Tooby.)

Krugman? Excuse me. He admits himself that he does not know much about evolutionary theory. He spends most of his talk noting the similarities between neoclassical economics and the Dawkins view of evolution, praising it. Except of course at the end he notes that it does not always work, and the messier, non-Dawkins stories are sometimes better. Duh.

Sure, there are some people who really dislike Gould (Dawkins, Dennett, and some others) who will make these statements about Gould not being respected by other evolutionary theorists. But his main idea, punctuated equilibrium, has entered the textbooks and is more studied and discussed than ever. Sorry, this is just whinging by the jealous or those who deeply disagree with him (Dawkins especially).

Oh, and as for me being taken in by bluffs, well, I have written on this stuff and have cited Williams, Hamilton, Maynard Smith, and numerous others, beyond those that Krugman claims are not cited by evolutionary economists, as well as Eldredge and Gould.

Finally, I have actually spoken with some of the most senior and distinguished evolutionary theorists, including some now dead. While Gould is often treated with some bemusement for his celebrity and eccentricities and the peculiar evolution of his views, I have not found these people viewing him as worthless or as having a rep worth mud. This is just silly bilge and drivel.

Finally, I would recommend you read Chapter 9 (I know, it is almost 300 pages long) of Gould's magnum opus. He does an excellent job of covering most of these controversies, with a fair amount of mea culpas for some of his own misstatements over the years. A lot more fair-minded than the junk in your post or in your links, frankly.

Hmm, well this is getting interesting from a layman's perspective. Time to crank up that expert detector with technical explanations...

Or perhaps it isn't worth your time? I won't understand it anyway. Sigh, back to my pre-cal homework.

In the meantime, there is some irony to be found in this link:

http://www.pandasthumb.org/archives/2005/10/on_the_beach_wi.html

Essentially Gould is misrepresented as a creationist by creationists! Or perhaps in the less extreme conclusion, he was agnostic about creationism.

Interesting assertions, Barkley; I'll check Chapter 9 when I get a chance. Meanwhile, I trust Tooby more than I trust you, to put it bluntly, to tell me what evolutionary theorists think of Gould. Of all Gould's critics, Tooby has directly earned my trust: The Psychological Foundations of Culture (Tooby and Cosmides) is one of my top five favorite science papers of all time and a marvel of precise thinking. Evolutionary psychology is perhaps the only field that demands and delivers even more careful thinking than evolutionary biology.

And every time I've ever encountered an idea of Gould's, it has been cloudy thinking dressed up with cloudy words. "Separate magisteria"? Give me a break. So I can see perfectly well for myself that he's running a bluff.

This post has generated some of the most passionate discussion yet on OB. This should be a warning sign: humans are especially prone to biases when it comes to closely cherished beliefs. Yet, I don't see a lot of hand wringing by either side about the possibility that the strongly held beliefs may be faulty.

Eliezer,

Gould makes his own mea culpas near the end of the chapter, so you can focus on that zone. Tooby is not an unreasonable guy, and it may well be that Gould's criticism of him was off-base. I have not read that particular exchange.

Let me put Gould in a slightly different frame. He was not the JK Galbraith of evolutionary theory, as Krugman bought into. He was probably closer to being the Milton Friedman, if on the opposite side of the political fence.

Friedman (like Dawkins also) was a great popularizer and author of best-selling books, like Gould and Galbraith. However, unlike Galbraith, he did win a Nobel Prize in economics, which most accept was deserved, even those who do not like Friedman's politics. He is also in all the intro texts, just like Gould, but unlike Galbraith. Even so, there are some, his worst critics, who have always been extremely critical of him, often in very hyperbolic terms and language. Also, he has been proven wrong about some things which he was closely associated with, notably hard core monetarism (he agreed last summer that central banks should not focus on money supply anymore, an ultimately reasonable guy, like Gould).

Nobels are not given out for evolutionary theory, although geneticists get them sometimes (Crick, Watson). So, Mayr, Simpson, Williams, Maynard Smith (he might have gotten an econ one), and other giants did not get one. If they were given, Gould would have gotten one. It is very simple. Whatever his many faults, and they were many, he came up with what has been the single biggest new idea (yes, one can find hints of it in Darwin and some others) in evolutionary theory of the last half century, punctuated equilibrium, now in all the textbooks, just like the most important ideas of Milton Friedman (and unlike Galbraith's major ideas). There are some who say it is wrong, but most say that it is at least partly right. So, Gould was very big and very important, even if he pissed a lot of people off.

Barkley Rosser: Punctuated Equilibrium is just placing an new buzzword on the common sense of of Gould's field, paleontology. Even if very broadly correct, it is not the single biggest idea in evolutionary theory of the last half century, not is it in the top ten. You don't find hints of it in Darwin and some others, you find it as essentially an implicit end of a continuum from completely uniform rates of change to completely discontinuous change. Most evolutionary biologists have assumed something closer to completely uniform change than Gould does, but he grossly exaggerates his differences from most biologists on this point.

Eliezer: As a general rule I would not recommend that you claim a paper or book as one of your all-time favorites if you last read it at a point in your life from which you have generally disowned your conclusions.

michael vassar,

Yes, but that "common sense" was widely ignored by most evolutionary theorists before Gould pointed it out. His approach is now in all the textbooks. He has also emphasized against some of his less fair critics that his original papers on this were specifically addressed to the paleontologists.

Another part of it, not so obvious and definitely not part of the general lexicon, is the idea of long periods of stasis. This was a point not at all emphasized by anybody prior to Eldredge and Gould, and was the real core of the original aspect of it.

So, what are the top ten ideas of the last half century ahead of it and where are they in the textbooks?

Regarding Darwin, the picture is muddled in that his remarks that look open to p.e. appear in later edtions of Origin of the Species, but not in the first one, where a more hardline gradualist scenario is presented.

Barkley,

Where would you put Gould on this scale:

(1) Someone who tries and succeeds in freeing their scientific work from their political biases. (2) Someone who tries but fails. (3) Someone who doesn't try. (4) Someone who not only doesn't try, but instead actively embraces their political biases and chooses their science appropriately with a goal of "improving society". (Obviously this approach only has value in fields such as some social sciences at the current time where absolutely disproving things is difficult).

It seems clear to me that Eliezer, Tooby, Smith, and Paul Gross all think Gould was (4), as do I. Further I find someone like this morally reprehensible, and I guess Eliezer from his strong language in this post does also. Now, putting aside whether Gould may have had one or two good ideas in his career, it seems to me from some of your comments here ("the peculiar evolution of his views", "oversimplifications or even obfuscations and confusions") that you actually don't disagree either. Yet you have no problem with this and actually seem to quite admire him on balance. Is this a fair comment? Why do you think there is this gap between yours and Eliezer's opinions?

I normally only read Robin's posts, but if watching Eliezer get so thoroughly told (Go Barkley!) can be assured to happen again with some frequency, I may have to change my policy.

This thread has more trolls than The Lord Of The Rings.

his eloquence was hardly unique among his competitors.

Are you sure he had any competitors?

Surely, his eloquence was not unique among similarly qualified academics, but probably few of them tried to get into the popular field. Maybe you need connections to get on the magazine circuit, but I suspect that the normal course of things are that the connections drag the scientist into writing. From there on, I imagine they're judged by their communication skills, not their content or connections. Also, Gould started when he was fairly young and energetic, as contrasted with, say, Lewis Thomas, who retired to writing when he was at an age Gould barely reached.

I find the response to Barkley Rosser's non sequitors disturbing. Sure, if he wants to pattern-match "gene's eye view" and start arguing about group selection, by all means argue about it, but don't accept his claim that it is relevant to this post! And punctuated equilibrium? Would it make a difference to any of this if Gould were an important theorist?

Tom: That isn't saying much - IIRC, LotR didn't even have that many trolls. The ones in The Hobbit were much more prominent.

Many, perhaps even most, of Gould's arguments were nonsense.

But many of the greatest minds in science have made nonsense arguments. We consider them great because they had at least a few ideas that were extremely worthwhile, because there was some gold in all the dross.

Rejecting Gould's positions on evolutionary theory merely because he also put forward the idiotic magisteria idea is dumb.

Ian Nowland,

I do not know where Gould fitted on that scale. I suspect he was in different places at different times.

OTOH, offhand I am not sure what is so "immoral" about #4, per se. Don't lots of people going into the hard sciences want to improve society, such as medical researchers? Certainly it is not surprising that people will think that what improves society is what agrees with their own ideology, even if perhaps you disapprove of their ideology. I think the potential problem here is the main topic of this blog: when people lie about or distort actual scientific findings or research in order to fit in with some ideological agenda. Certainly people have charged Gould with being motivated for parts of his research by his ideology. However, I am not aware of anybody successfully claiming that he actually distorted or lied about findings in the process of doing so. If not, then the charge of "immorality" is way overblown, just like most of Eliezer's charges against him, drawn from sources who were his worst enemies and strongest opponents.

Douglas Knight,

Not sure without further discussion what my "non sequitors" were, but your statement about Gould as a (non) theorist is amazingly empty. How is that you say hs is not an important theorist? While some have labeled it as "just relabeling," or something like that, I have yet to see anybody here offer up even one, much less ten, new ideas in evolutionary theory more important than Gould's of punctuated equilibrium.

I will offer one: coevolution, due to Paul Ehrlich. However, even more than punctuated equilibrium, this is one that one can find strong precursors of in Darwin, big time.

Progress is not the same thing as complexity increase. While I agree that there can be upper bounds on complexity increase in evolution, this doesn't really have that much relevance to the question of whether evolution has any cumulative direction. The latter is something I consider to be an open question.

As any engineer knows, great increases in capability often occur through the removal of complexity, not its addition. This happens more frequently than you'd think. It's part of why, for instance, computers have gotten better, faster, and cheaper. If you look at a computer today you'll see that it has less components than a computer did a decade or so ago. Evolution can of course do the same thing. Fitness can increase through loss of complexity. The simplest and most straightforward example would be the knockout of a detrimental gene, while a more complex example would be the construction of a complex genetic framework or "scaffolding" followed by the emergence of a simpler system within that framework followed by the dissolution of the framework's no-longer-necessary parts.

So while Williams' work may show the existence of upper bounds on complexity, it does not disprove the idea that there might be some kind of directionality that could be called progress in evolution. If it exists, it seems to be something that we haven't learned how to quantify or explain quite yet.

Barkley Rosser,

oops, I meant "if...weren't."

Your invocation of group selection was completely irrelevant. It's so irrelevant, I refuse to elaborate. I find it hard to believe you continued reading past the phrase "Williams revolution." It's worth pointing out that Tooby explicitly praises Gould's promotion of group selection.

Punctuated equilibrium was an ad hominem response to an ad hominem attack, so it's not irrelevant, but it's pretty tenuously connected.

The problem isn't that Gould was wrong or over-promoted his own ideas; the problem is that he massively misrepresented the state of the field. The problem is in his popular writing, not his professional writing (though Maynard Smith does mildly criticize them as well).

Yeah, what Adam Ierymenko said:-) about hitting a complexity limit being not at all synonymous with stopping progress. Except that I was going to say "computer programmers" instead of "engineers", and I was going to use the example that when duplicate functionality in the mitochondrial genome and main-cell genome gets replaced by shared functionality, the organism tends to win back some ground from the the Williams limit you described. And, incidentally, the mitochondrial example is very closely analogous to something that practicing computer programmers pay a lot of attention to: Google for "once and only once" or "OAOO" to see endless discussion.

It's vitally important when spreading a new scientific idea to come up with a catchy name for it. "Punctuated equilibrium" was very good. However, the idea behind the name was very very fuzzy. It was a grab-bag of ideas, that never quite fit together. Where did the "hopeful monster"s fit in? It turns out they didn't. Punctuated equilibrium did not make sense in any unified way though a collection of disparate ideas were supposed to be it.

I finally found the concepts expressed clearly in a book by Eldredge. All the various obvious explanations were included. There was nothing mystical. There was nothing new. Standard population ecology concepts fit the whole thing. Unless of course Gould was describing other things that Eldredge didn't consider.

The spandrels concept was like something Eliezer might come up with. A fine name for a simple concept about how people sometimes misunderstand things. The "trait" that you notice may be a side-effect of what's selected. It's worth reminding beginners about that and worth remembering. It isn't more special than the average of Eliezer's hundreds of bias ideas.

Gould talked as if he had something that standard science did not explain easily. He was misleading. No hopeful monsters were needed. Everything he described was implied directly from standard approaches. Gould mostly did not describe solutions to his conundrums -- I think because if he had, he would have made it obvious that he was not actually posing any problems to be solved.

"I have yet to see anybody here offer up even one, much less ten, new ideas in evolutionary theory more important than Gould's of punctuated equilibrium."

Can you describe just what the dea is? When people ask for a description of Gause's Law or Weber's law or Fisher's Fundamental Theorem etc you could say pretty clearly what the idea is, right?

Gause's Law: No two species living in the same environment will live quite the same way. They will tend to differ in ways that are critical to their survival, because if they were in direct head-to-head competition for the same limiting factor one of them would tend to win and the other would go extinct. (It follows that two species that too much share the same critical limiting factor will tend to evolve in separate directions to reduce the overlap. They will evolve toward different ecological niches.)

Weber's Law: Animal senses distinguish differences, and the differences they can distinguish tend to be some fraction of the size of the stimulus. So if you can tell a 1% difference in how bright a light is when it's very dim, you can likely tell a 1% difference in brightness when the light is very bright also. If you can tell a 1% difference in sound volume for faint sounds it's likely to be around 1% for loud sounds too. This all happens within limits and it doesn't always work, but it's the way to bet.

Fisher's Fundamental Theorem: The rate of natural selection is directly proportional to the variance in fitness within the populatoin. This follows directly from the usual definitions of fitness and natural selection. If all individuals are just as fit then there's no selection, if there's a big difference in fitness then the less-fit are removed faster than with a small difference.

Can you describe what Punctuated equilibrium says? Perhaps the 2nd-most important new idea in evolutionary theory, can you describe the idea clearly?

Kaj: By jingo you may be right. But the question is, if the population of trolls is falling between The Hobbit and LoTR, is this an example of species-level selection? Because I can see that coming in handy. Imagine:

Johnny Creationist: "The Bible says God created the Earth in six days!"

Me: "Well the Lord of The Rings says we evolved by punctuated equilibrium!"

Johnny Creationist: "Damn! Your logic is unassailable! Well, I'm an atheist now."

shake hands

"You are in way over your head on this one. It is clear that you are behind on current literature. The "selfish gene" is old hat and out of date. Multi-level evolution is increasingly widely accepted by many geneticists, with such mechanisms as reciprocal altruism being keys."

Perhaps you shouldn't be so hasty at pointing the "over your head" finger at others. Reciprocal altruism is NOT about multilevel selection. It's good old individual selection - helping others on the assumption that the help will later be returned. This explanation assumes a fitness benefit (on average) for each participating individual, invoking group selection here is not necessary.

Douglas,

I am perfectly willing to buy that Gould's scientific writings are more defensible than his popular writings.

J Thomas,

In my later comment I did not repeat my earlier comment. There I had specified, "one of the top ten ideas in the last half century," not of all time. So, Weber's Law dates from the 19th century, and those of Gause and Fisher from the 1930s. Not relevant. Still waiting for someone to come up with something other than coevolutions, which, as I have already pointed out, is definitely in Darwin, if not the term, neologized by Ehrlich just as Gould and Eldredge neologized punctuated equilibrium, whereas the latter is only ephemerally in Darwin, at best, and only then in later editions of Origin of the Species.

A definition? That a substantial amount of speciation occurred during (geologically) relatively short periods of time, between which were long periods of relative stasis without actual speciation, although gradual changes were always occurring. Gould himself in TSOET emphasizes that p.e is not in contradiction with the neo-Darwinian synthesis of the 1930s and 40s. And I know from talking to some of the developers of that theory that they do not disagree with that statement of his. Rather the issue is seeing this particular outcome as widespread and important rather than as some weird sideshow not even openly discussed, much less labeled. Sorry, but Gould and Eldredge did in fact make a theoretical breakthrough of great importance. Heck, Einstein himself argued that relativity was already implicit in the work of Galileo.

windy,

You are correct about reciprocal altruism. But that does not undo the more general argument. Apologies for getting sloppy, but indeed plenty of people are taking multi-level evolution seriously. And, more to the point, it remains the case the Eliezer is simply dead wrong about the consensus of evolutionary theorists regarding the status of Gould. Again, one might as well make claims about the historical significance of Milton Friedman based on the opinions of Marxist economists. That is effectively the equivalent of what Eliezer attempted here in this post.

BTW, creationists did use Gould at a certain point based on his rhetoric criticizing "fundamentalist Darwinism" to dump on Darwinism and evolutionary theory in general. This probably did feed into the extreme annoyance by some evolutionary theorists with Gould at one point.

"A definition? That a substantial amount of speciation occurred during (geologically) relatively short periods of time, between which were long periods of relative stasis without actual speciation, although gradual changes were always occurring."

That's it? But we've known about adaptive radiation for a very long time. What's the new idea here, beyond this obvious observation?

I didn't quote those three to say they were important new ideas, I quoted them to give an idea what sort of explanation I'd accept. Reasonably short, showing what it was about, leaving out caveats and exceptions etc.

You haven't begun to show what Punctuated Equilibrium was about. I don't think it can be explained simply because there isn't a central new idea there.

Before deciding that Gould's theory was wrong or unimportant, read something from 2007. www.biology-direct.com/content/2/1/21

Douglas, I don't claim that Gould's theory was wrong or unimportant. I claim that Gould's theory was incoherent to the point that there's no way to tell whether it was wrong or unimportant.

It's like deciding whether a horoscope or a Rorschach test is wrong or unimportant.

Eldredge's theory of Punctuated Equilibrium was unimportant. Eldredge explained it well enough to see what he was saying. Gould apparently was talking about something else.

J Thomas,

Well, this is about to scroll off, but I would ask: 1) Did anybody prior to Eldredge and Gould point out this argument? (Answer: NO) 2) Is it important? (Answer: YES, at least it is in all the textbooks and is widely studied and discussed) 3) Do you or anybody else have a more important new idea in evolutionary theory that has appeared in the last half century??? I asked this some time ago. Nobody has stepped forward, other than myself with coevolution. Are you going to? Are able to? I doubt it.

You are just making yourself look as foolish and off the wall on this matter as Eliezer.

Barkley, you have given us no idea what idea it is you say is so important.

Is your claim that what it means is that there are some times when evolution of bony parts is fast, and other times when it is slow? That follows directly from "adaptive radiation", which has been known for a very long time and which suggests a reason. Nothing new there.

If it's such an important idea surely you can tell us what it means. Take all the time you want. Don't worry about it scrolling off, we'll wait for you.

At this point the argument isn't whether PE is important. The question is whether you know what it says. We can talk about how important it is after you show us you know what it means.

J Thomas,

I have already given a definition, which is close to what you will find in Wikipedia, if you go look there. Of course one can argue endlessly (and many do) about whether or not particular cases fit the definition or not.

Adaptive radiation can be viewed as a special case of punctuated equilibrium, although it is one that involves genetic drift, first identified by Sewall Wright in the 1930s, with Mayr emphasizing geographic separation arising from genetic drift as a key form of this. Adaptive radiation in particular refers to a case where a population wanders into an isolated and underpopulated area, where it can "radiate" quickly into a bunch of different niches. The classic case goes all the way back to Darwin, his finches on the Galapagos islands, although the term was cooked up much more recently. But, this is only one way one can get such rapid evolution in a short time.

The crucial figure on genetic drift was Sewall Wright, one of the "trinity" of the 1930s neo-Darwinians, along with Fisher, whom you praised, and Haldane. Wright's position with them is very controversial. He lived to 1988, and I actually knew him and talked with him on several occasions. Gould's account of his interactions with Wright in TSOET is fascinating, a very complicated relationship. I also note that Wright's statistical work had importance for econometrics, with a paper by him and his father being the first to identify the identification problem.

Anyway, J Thomas, I think you are the one who needs to show that you know what the hell you are talking about. No, you do not control when this scrolls off so that "we are waiting." We are waiting, or at least I am, for you or anybody else here to come up with an idea, any idea, that you can claim on any grounds, you describe them, please, more important in evolutionary theory in the last half century. Neither you nor anybody else has done so yet, despite my repeated challenges.

BTW, I have a website, easily found as the second entry by googling my last name. You can go read my own publications on these matters. Have you ever published anything on evolutionary theory? Has Eliezer besides on blogs?

"A definition? That a substantial amount of speciation occurred during (geologically) relatively short periods of time, between which were long periods of relative stasis without actual speciation, although gradual changes were always occurring."

Mr. Rosser, This definition is very much like that in Wikipedia, true. As stated, it follows directly from observed adaptive radiation -- which has had explanations involving genetic drift. There is nothing new in this definition. It is not a new observation. Tnere is no new idea expressed here beyond the old observation. This is a pretty shiny packaged christmas present with nothing in it.

Can you state a definition that shows what's special about punctuated equilibrium? If you can't, then there's no reason to think you understand it.

I looked at your economics homepage. I think you're studying a fascinating specialty and I wouldn't be at all surprised if I learn interesting and useful things as I read your publications. Thank you for the link.

J Thomas,

So, we are running in circles. Adaptive radiation is just a special case of the more general phenomenon, although it might not be if the rate of change/speciation continues rapidly without convergence to some equilibrium. Adaptive radiation was described in Darwin, without the term, and in some later editions he did note the variabilty of rates of evolution, although he did not exress the idea that much speciation would occur very rapidly followed by long periods of substantial stasis. This idea is never stated anywhere in Origin of the Species, nor does it appear in any writings prior to Eldredge and Gould.

One could say that what is involved here is the stating of a hypothesis, that is then given a label. It is a much broader idea than just adaptive radiation arising from genetic drift, which is indeed something that had been described earlier, but I repeat is only a special case and does not include the crucial addendum regarding long periods of stasis. Indeed, as I noted earlier, it is this latter point that is really the new part of E and G's idea of p.e. Certainly that speciation might occur rapidly under various circumstances, especially when the general environment is changing rapidly, as for example during a Cuvierian "catastrophe" such as wiped out the dinosaurs, was noted previously. But such catastrophes are also essentially special cases, just as is adaptive radiation. It is the adding of the long periods of stasis that distinguishes the idea/hypothesis, or whatever you want to call it. But let us be clear "adaptive radiatiom" does not equal "punctuated equilibrium."

BTW, you will not find my most extensive discussions of evolutionary theory on my website. The most extended discussion appears in my 1991 book, From Catastrophe to Chaos: A General Theory of Economic Discontinuities, Kluwer, especially in Chap. 12.

Again, probably for the last time, I await from anybody an idea appearing in the last 50 years, besides coevolution, that has been more important, or perhaps more influential, than punctuated equilibrium. Without such an offering, I would say my case rests, and very solidly.

I don't know exactly when things were invented, but just ten off the top of my head: what about intragenomic competition, epigenetics, Price's Equation, reciprocal altruism, the gene's-eye-view, multilevel selection, adapted enforcement of boundaries leading to the emergence of cells and multicellular organisms, sociobiology, gene regulatory networks, and the molecular clock as a means of tracking adaptive pressures? And before you say that any of these things were preinvented before 1957, I would like to know if an equally strong argument could be made for the preinvention of punctuated equilibrium.

I first read about adaptive radiation in 1965 or so, I believe it was in Ralph Buchbaum's Basic Ecology. Or maybe it was World Book encyclopedia. It said that we get lots of speciation events right after a big extinction event, and then they slow down for a long time. You say that this is a special case. Because somebody hypothesised that it had something to do with genetic drift, while nobody hypothesised that PE had anything to do with genetic drift?

There is nothing you have described about PE that isn't true about adaptive radiation. Can you say what's special about PE? The fossil record already showed us that we got long periods of stasis in hard-parts, that was not a new observation. Did Gould perhaps suggest a new reason for the stasis, one that the population ecologists hadn't already stated?

I could easily be wrong that there's nothing here that's important and new. But you have said nothing to show what that something might be, or why it's important. If it's one of the fundamentally important things, why can't you say simply what's new and important about it?

Is it actually such a complicated idea that you can't explain the fundamentals in a short post?

Triver's explanation why there is so much self-deception. Parent-offspring conflict.

I'd also like to request an explanation of why punctuated equilibrium matters to anyone other than paleontologists. So rates of morphological change per se are sometimes fast and sometimes slow. (I believe modern research has strongly suggested that the "equilibrium" part of PE actually exhibits equally strong selective effects and equal rates of genetic change, just not differently shaped organisms.) So what? So the old gradualists are mistaken? Okay, but then what? What follows from this?

Contrast to, say, sociobiology which has reorganized our understanding of every social animal species. Contrast to reciprocal altruism which has spawned a new field of evolutionary science as well as game theory. Contrast to intragenomic competition which is opening up whole new vistas of explanation for vast redundant stretches of DNA, and showing how the existence of a multicellular organism is a constant battle to protect the fairness of the reproductive organs. Contrast to the gene's-eye-view revolution that destroyed hundreds if not thousands of specific biological misconceptions about the purposes of adaptations. These revolutions reshaped our anticipations, not just our beliefs.

What is punctuated equilibrium, that it should even be mentioned in the same breath as such innovations? And what is Gould's specific contribution to the concept, besides the name? Enlighten me, if I am ignorant.

The prediction and discovery of segregation distortion genes looks more important to me than PE, given that PE looks like nothing new or important at all.

The discovery of transposible elements may have happened in 1950 but the implications for population genetics had still not been examined the last time I looked. They have the potential to revise most of the early genetics conclusions. For example, Fisher and Wright had an argument about evolution of dominance which Wright won. It would take longer than the lifetime of an average species to select for a gene that affected the dominance of another gene, given the assumptions of the time. But that was assuming genes that were evolved to affect dominance that arose in place by mutation and natural selection. A transposon that can move around in the genome, causing the block of DNA it inserts into to become dominant or recessive, could be selected very fast. A copy that inserts in a gene that is favorably selected and makes it dominant would increase in frequency with the gene it improved. And a copy that happened to insert into a gene that is deleterious and makes it recessive will at least survive much longer than it would otherwise, maybe long enough to transpose copies elsewhere. If they switch between making their genes dominant and recessive at a fairly low rate -- say 10^-4 -- they won't be subject to much unfavorable selection when that's in the wrong direction but they'll be nicely selected when it works.

Many of the assumptions of the classical theoretical geneticists are wrong given the existence of transposible elements. The whole thing has to be thought out fresh.

Should we measure the importance of the ideas discussed by citations?

Not sure that's fair in this case, even if we restrict to scientific literature; I've read a number of citations of Gould, but usually along the lines of "Contrary to Gould and Lewontin (whenever), there is no reason to think XYZ is a spandrel," etc.

Ah, finally got some real action out of you guys. Not bad.

Eliezer,

Pretty decent list. Of course, no way we are going to resolve this, especially if we are going to rule out such "objective" measures as citations, where indeed I suspect p.e. will beat most of those. Also, there is the complication that several of those are really extensions of each other, such as Price's equation providing conditions for multi-level evolution, which is also partly connected to reciprocal altruism, not to mention that the since the gene's-eye view tends to imply gradualism, which is hardly dead, p.e. challenges it.

J Thomas,

I would agree that it is of the greatest importance to palenotologists. One importance of p.e., ironically, is that although creationists sometimes tried to use Gould's arguments against evolution, p.e. provides an answer to the old creationist gibe about "where are all the missing links?" (although somehow those folks conveniently ignore good old Archeopteryx somehow).

We still haven't heard anything about what PE says that's new or interesting. I can't agree that arguments against creationists is that big a deal.

OK, it could be somehow important to paleontologists. But paleontologists have known all along that skeletal morphology is mostly fixed. After all, a species that has particularly variable morphology is likely to be classified as multiple species. And so the dog that didn't bark in the night is that when you look at the bones from one species they seldom show slow gradual directional change over the lifetime of the species. Who would have predicted that they would?

Here is a simple ascii art story.

When a mutation spreads through the population through selection, you can usually expect it to increase logisticly.

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How would this graph look if you took only a few samples for each timepoint and compressed the scale?

|.......**
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There's nothing profound about this.

Jesus H Xrist this debate has devolved (not an evolutionary term, I know, but perhaps a literary-dialectical one). I've been out of commission for a while and just found it. I would offer some of my views, but I don't consider a comments-section on a blog to be conducive to effective discourse. Why doesn't each party take some time to formulate a more formal argument against the other?

The only thing you have to gain is the a better map of the territory.

Finally, I would recommend you read Chapter 9 (I know, it is almost 300 pages long) of Gould's magnum opus. He does an excellent job of covering most of these controversies, with a fair amount of mea culpas for some of his own misstatements over the years. A lot more fair-minded than the junk in your post or in your links, frankly.

Well,

I got the book out of the library,

Managed to quickly flip through Chapter 9 tonight, after finishing a rather long day's work,

(Mostly because the book is due back at the library tomorrow),

And I've got to say, Gould's reputation as a clear writer is overrated. I didn't find any exonerating evidence for Gould in the parts of Chapter 9 that I quickly flipped through. It's difficult to read a book you can't trust -

- as exemplified, for example, in the very Table of Contents, where in Chapter 7 on "The Modern Synthesis as a Limited Consensus", we see Gould (and his book was published in 2002), discussing "Synthesis as Hardening" and "The Later Goal of Exalting Selection's Power", discussing books published in 1937, 1951, 1944, 1953, 1942, and 1963. In short, the huge revolution that started in 1966 doesn't seem to exist so far as Gould is concerned. There was a subsection of Chapter 7 labeled "Adaptation and Natural Selection", which I flipped to, but it's not about Williams's book at all, just a random section with the same name!

Lamarck gets a mention in Gould's Table of Contents. So does William Paley. Not George Williams, though. Apparently it's harder to hit a moving target.

Going to the index, I did find a number of mentions of George Williams. I flipped to a discussion of "adaptation and natural selection" on pages 550-554. The discussion is about group selection. Then Gould starts going on about some mysterious sin called "reductionism", of which Williams is apparently guilty, but what kind of false experimental predictions result is never quite explained, at least not in the parts that I read.

In short, if Gould departed from his policy of railing against the romanticism of the pre-Williams period as if they were current beliefs that Gould alone had dispatched - this being his great sin bordering on warped plagiarism - his redemption is not visible from the small parts of this gigantic book which I was able to peruse, before I returned the tome to my library.

EO Wilson has a section in his autobiography, /Naturalist/, on what Gould and Lewontin did after the publication of Wilson's /Sociobiology/. They formed a study group, which met every week to criticize Sociobiology, then after a few months, published their results.

The kicker is that they held their meetings about a 30-second walk from Wilson's office in Harvard - but never told him about them.

This proves to me that science and truth never were their primary concern.

"academic debates are personally brutal, not because the stakes are so large, but precisely because the stakes are so small." - Anonymous

Isn't the byline usually given as "Stephen Jay Gould"?

I realize this discussion is a few years old, but I just came across this post while browsing through the sequences, and I wanted to put in a word for Gould's book "Full House" that was the main target of this post, since I just read it last year.

First of all, only a third of the book is about evolutionary biology at all. The part I remember more was a discussion of the disappearance of .400 hitting in baseball, using similar statistical arguments.

Second, in the the section that was about evolution, I did not come away with the impression that he was implying that most evolutionary biologists believed in increasing complexity, nor that he was setting himself up as a hero. As I remember it, the tone was much more "many laymen have this impression, because many biology textbooks and other references outside the field tend to depict evolution as a pyramid or progression, with more complex organisms at the top."

And you know what? He was right. I'm a layman, and I did have that general impression, and it was useful to read a detailed explanation of why it's wrong. And I'm also a baseball fan, and I had not thought of his argument about .400 hitting before I read it, and I found it pretty interesting.

Overall I found him slightly pompous as a writer but nothing like the way Eli (or others like Dawkins) have described him. I've never read any of his other books, and I'm absolutely not qualified to comment on the debate in the comments about the merits of PE or his scientific stature. But I think the content of "Full House" is not described accurately in the post.

This is several years old, as Peter noted, but I wonder what the reaction to Gould is now, in light of the recent publication in PLoS Biology demonstrating that in his Mismeasure of Man, Gould manipulated the data until he got the result he wanted. http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1001071 Essentially, he made it up. In the process, he has deliberately deceived thousands or millions of readers, he has tarnished the reputation of a scientist (Morton) and stymied significant research (in sociobiology, for example). Why did he do this? What was his personal agenda? And why are people who should know better defending him here?

I'm going to try and do this formally once I've got more information theory under my belt, but I suspect that a very large portion of the "deaths" needed to sustain information in a population of large animals actually occur in gametes and embryos. Both human eggs and sperm for instance undergo a competition process that selects the best, and pre-implantation lethality in embryos is extremely common - which is one of the reasons for humans' very low fertility rate.

Some points. Sorry to not give references. I hope later to write up a lot of this stuff, but meanwhile I think a warning on Yudkowsky's warning is desirable asap.

• From memory, Jerry Coyne is actually a fan of Gould - don't know where Tooby thought that he wasn't. Ditto David Jablonski.

• Gould 'finished' his 1400 page plus magnum opus, The Structure of Evolutionary Theory, only a few months before he died of cancer. Yes, it is often woefully verbose, with repetitions and overlaps that would otherwise be imho inexcusable. And his various thoughts on different levels of selection are not well structured (but imvho these are quite difficult for non-mathematicians / programmers to think about - it takes a bit of practice before you think about them clearly). But there is gold buried in his sea of words.

• Gould was fighting, inter alia, against biologists' inertia re 2nd order selection, and against Dawkin's earlier mistake (since recanted) re the Selfish Gene. See, e.g. Darwin in the Genome by Lynn Helena Caporale for a hint of what Gould was up against. And Mayr changed his mind during his long career re speciations. I groan at some of Gould's overstatement, but sorry Yudkowsky, at least on this topic, your overstatement is worse than Gould's.

• I haven't read all Barkley__Rosser's comments, but at least the first few make a lot of sense to me - nice work Barkley.

Note that the "Williams revolution" that Gould is widely seen as having stolen (and which Tooby accuses him of having stolen) is not what Eliezer Yudkowsky says it was. It had nothing to do with the argument in Full House, nor to do with Williams' equilibrium-amount-of-information argument, which is not at all the most widely-remembered part of Adaptation and Natural Selection. (I actually think Gould's argument makes sense, or partial sense, insofar as it's a response to the lay public's impression of the big-scale evolutionary history of life on Earth as having been driven to climb a ladder from simplicity to complexity. Why did fish evolve after single-celled organisms?—Well, it could hardly have happened the other way around, could it?)

The revolution in question is about criticisms of pan-adaptationism ("panglossianism"; "ultra-Darwinism"), i.e. the habit of assuming every trait you can think of in an organism was naturally selected for some purpose [and the habit of speculating what that purpose was]. (BAHfest, the "festival of bad ad-hoc hypotheses", invented by SMBC, also mocks that habit, which is still around despite the best evolutionary biologists' best efforts; e.g. I was in a first-year bio class where the TA tried to illustrate selection thinking by speculating that caterpillar mouths open sideways so they can walk along the edges of leaves and eat; sure, maybe, but maybe it's relevant that insect mouths in general do that, and maybe vertical vs. horizontal makes little fitness difference anyway.)

Williams gave the good example of a flying fish returning to the water. Is it fitter to eventually fall back into to the water than to not? Probably, sure. Does that mean it's an adaptation? No, of course not, it's just the default you should expect because of gravity. Indeed the first sentence in Adaptation and Natural Selection was (1966):

Evolutionary adaptation is a special and onerous concept that should not be used unnecessarily, and an effect should not be called a function unless it is clearly produced by design and not by chance.

Gould and Lewontin also criticized pan-adaptationism in their paper, "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme" (1979). They gave a memorable metaphor, and coined the term "spandrel" for traits which are not themselves adaptations, but developmental side-effects of other traits (e.g. male nipples, and female orgasm, in mammals). The metaphor came from architecture: the spandrel (or pendentive) is the tapered triangular bit that results, if you put a dome on top of rounded arches; the spandrels in St. Mark's Basilica have such beautiful mosaics that a naive tourist might assume the spandrels were put there for the mosaics; but they'd be wrong. I agree Gould and Lewontin should have cited Williams, but I'm glad they gave us the metaphor; it's almost as beautiful as the spandrels themselves. They also coined the usage of "just-so stories" (after Rudyard Kipling) as an insult-term for untestable post-hoc speculation.

The context of Tooby's letter to the editor is that Gould had been accusing evolutionary psychologists of being Panglossian just-so storytellers. Tooby was replying that they knew perfectly well not to be Panglossian; Williams was their doyen after all; that they were in the business of testing hypotheses' predictions, not just speculating about the past; etc.

Anyway, as for Gould himself—I wouldn't say never read him, but I would say read him with a big shot of salt; and yes, if he implies he's revolutionizing something, don't believe him. As Maynard Smith says, his essays are often excellent. But e.g. he way-overblew how big a deal punctuated equilibrium would be even if true (see the chapter about it in Richard Dawkins' The Blind Watchmaker). And so on.