Followup to:  Natural Selection's Speed Limit and Complexity Bound

If you've read anything Stephen J. Gould has ever said about evolutionary biology, I have some bad news for you.  In the field of evolutionary biology at large, Gould's reputation is mud.  Not because he was wrong.  Many honest scientists have made honest mistakes.  What Gould did was much worse, involving deliberate misrepresentation of science.

In his 1996 book Full House: The Spread of Excellence from Plato to Darwin, Stephen J. Gould explains how modern evolutionary biology is very naive about evolutionary progress.  Foolish evolutionary biologists, says Gould, believe that evolution has a preferred tendency toward progress and the accumulation of complexity.  But of course - Gould kindly explains - this is simply a statistical illusion, bolstered by the tendency to cite hand-picked sequences like bacteria, fern, dinosaurs, dog, man.  You could equally well explain this apparent progress by supposing that evolution is undergoing a random walk, sometimes losing complexity and sometimes gaining it.  If so, Gould says, there will be a left bound, a minimum at zero complexity, but no right bound, and the most complex organisms will seem to grow more complex over time.  Even though it's really just a random walk with no preference in either direction, the distribution widens and the tail gets longer.

What romantics, ha ha, those silly evolutionary biologists, believing in progress!  It's a good thing we had a statistically sophisticated thinker like Stephen J. Gould to keep their misconceptions from infecting the general public.  Indeed, Stephen J. Gould was a hero - a martyr - because evolutionary biologists don't like it when you challenge their romantic preconceptions, and they persecuted him.  Or so Gould represented himself to the public.

There's just one problem:  It's extremely unlikely that any modern evolutionary theorist, however much a romantic, would believe that evolution was accumulating complexity.

There was once a time when many evolutionary biologists had a romantic conception of progress, evolution climbing ever-higher mountains of complexity, dinosaur to dog to man.  And there was a hero who challenged that widespread misconception.  The hero was George Williams, his challenge was successful, and his reputation rests securely in evolutionary biology today.

In a population at equilibrium, harmful mutations will be eliminated by death (or failure to reproduce) at the same rate they are introduced by copying errors.  A very severe mutation may be eliminated by an embryo that fails to develop, but a mutation that's lethal only one time out of 10,000 may spread to 10,000 people before it starts to be eliminated.  It takes the same amount of selection pressure to support minor or major adaptations; whether the adaptation was a big one or a small one, at equilibrium, mutations must be eliminated at the same rate they are introduced by copying errors.

A population cannot sustain too high a selection pressure - too many deaths or failures to reproduce - without dying out.  And it requires the same amount of selection to support any given amount of DNA against the degenerative pressure of copying errors.  This, in turn, implies an upper bound on the amount of DNA that can be sustained by selection against the degenerative pressure of copying errors.

The upshot, as George Williams wrote:

A certain amount of information is added by selection every generation.  At the same time, a certain amount is subtracted by randomizing processes.  The more information is already stored, the more would mutation and other random forces reduce it in a given time interval.  It is reasonable to suppose that there would be a maximum level of information content that could be maintained by selection in opposition to randomizing forces...

The view suggested here is that all organisms of above a certain low level organization - perhaps that of the simpler invertebrates - and beyond a certain geological period - perhaps the Cambrian - may have much the same amounts of [meaningful] information in their nuclei.

Saying this did not make Williams a heroic, persecuted martyr.  He simply won.  His arguments were accepted and biology moved on.  The book quoted above is Adaptation and Natural Selection, now showing its age but still considered a great classic.  The shift to a gene's-eye-view in evolutionary theory is sometimes called the "Williams Revolution", the other founders being Hamilton, John Maynard Smith, Trivers, and Dawkins as popularizer.  In short, Williams was not exactly Mr. Obscure.

And Williams wrote in 1966, thirty years before Gould wrote Full House.

If Gould had simply stolen Williams's ideas and presented them as his own, then he would have been guilty of plagiarism.  And yet at least the general public would have been accurately informed; in that sense, less damage would have been done to the public understanding of science.

But Gould's actual conduct was much stranger.  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.  Then Gould presented his own counterarguments to these no-longer-advocated views, and they were bizarre.  Evolution is a random walk in complexity, with a minimum at zero complexity and no upper bound?  But there is an upper bound!  Sheer chance explains why dogs are more complex than dinosaurs?  But they probably aren't!

Why did Gould behave thus?  Two observations:  One, to bring order to a scientific field, it must first be in chaos.  Two, plagiarism is a crime that everyone understands.

Gould undid the last thirty years of progress in his depiction of the field he was criticizing, pretending that evolutionary theory was in chaos, so he could depict himself as heroically bringing order to it.  If Gould had also redid the accepted solutions as his own, he would have been caught, tried, and cast out of the public eye.  Newspaper editors may not be interested in mathematical arguments about evolutionary biology, but they understand stories about plagiarism and theft.  Once Gould's copying had been laid out next to the original, and eminent scientists attested to the identity, it would have been over.

So instead Gould committed a crime so bizarre that it couldn't be explained to editors.  He stole Williams's chaos.

(Incidentally, Gould's notion of a random walk in complexity has the same quality as the rest of his argument.  A genome acquires a beneficial allele at a readily calculable speed and probability, and until the complexity reaches equilibrium, new adaptations will tend to be acquired faster than old adaptations are lost to copying errors or environmental shifts.  The fewer adaptations have been acquired by a genome, the fewer are likely to be lost to a given event.  If complexity starts far below the equilibrium level, it will tend to increase.)

All this that I have said, was a common pattern throughout Gould's "work".  And all this that I have said, is no news to professional biologists.  Here's John Maynard Smith:

"Gould occupies a rather curious position, particularly on his side of the Atlantic. Because of the excellence of his essays, he has come to be seen by non-biologists as the preeminent evolutionary theorist. In contrast, the evolutionary biologists with whom I have discussed his work tend to see him as a man whose ideas are so confused as to be hardly worth bothering with, but as one who should not be publicly criticized because he is at least on our side against the creationists.  All this would not matter, were it not that he is giving non-biologists a largely false picture of the state of evolutionary theory."

John Maynard Smith was a genuinely great evolutionary biologist, the sort of man that Gould pretended to be.  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. 

I am not calling Gould a scoundrel because he was wrong; honest scientists can make honest mistakes.  But Gould systematically misrepresented what other scientists thought; he deluded the public as to what evolutionary biologists were thinking.

It is as if someone presented geocentric epicycles as the current belief in 21st-century astronomy, sharply criticized the complexity of all those circles orbiting circles, and argued for their own simpler model of planets that move in straight lines.

Did Gould deliberately lie?  If not, he executed one of the most epic feats of self-deception in the history of marketing.  The eminent John Tooby speaks:

"Although Gould characterizes his critics as "anonymous" and "a tiny coterie," nearly every major evolutionary biologist of our era has weighed in in a vain attempt to correct the tangle of confusions that the higher profile Gould has inundated the intellectual world with.  The point is not that Gould is the object of some criticism -- so properly are we all -- it is that his reputation as a credible and balanced authority about evolutionary biology is non-existent among those who are in a professional position to know...
These [major evolutionary biologists] include Ernst Mayr, John Maynard Smith, George Williams, Bill Hamilton, Richard Dawkins, E.O. Wilson, Tim Clutton-Brock, Paul Harvey, Brian Charlesworth, Jerry Coyne, Robert Trivers, John Alcock, Randy Thornhill, and many others."

If Gould, after receiving that many corrections, managed to still not know the actually current beliefs in evolutionary biology, he must have had neutronium earplugs.  I'm not saying it's impossible, though, because it's amazing what people can not-know when their reputation depends on it.  But there comes a point in self-deception where it becomes morally indistinguishable from lying.  Consistently self-serving scientific "error", in the face of repeated correction and without informing others of the criticism, blends over into scientific fraud.

And after all this, Gould is widely believed, by the general public and even by many scientists outside evolutionary biology, to be an evolutionary theorist of honorable reputation!  It is as if Immanuel Velikovsky had managed to make himself into the public face of astronomy.

If you have read one of Gould's books, you are not to blame; but you must now do your best to un-believe it all - especially all the implied beliefs in evolutionary biology that Gould seemed to be attacking.

And so as not to be accused of plagiarism myself, many others have said much of what I said here - only in politer academic language, with longer sentences, and without that specific example.  I thought it deserved a sharper presentation, for the benefit of the popular audience that Gould deluded; and a clear-cut example of Gould's "work", to show what the fuss is about.  Many academic writers on Gould could not speak as sharply as Gould deserved.  As I have no fear for my own reputation, I will say it plainly:  One way or another, knowingly or unknowingly, Gould deceived the trusting public and committed the moral equivalent of deliberate scientific fraud.


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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... (read more)

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 o... (read more)

"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 ... (read more)

We may also have better error-correcting mechanisms than those simple invertebrates do, though I'm not enough of a biochemist to know myself. (By the way: "invertebrate" is a terrible category---akin to being a non-unicorn---and gets especially bad when the concern is intelligence; it ranges from the near-mindless oyster to the brilliantly clever octopus.)
Good point. Can you (or anyone else) suggest anything better?
Don't try to refer to non-chordates as a single class, would be my suggestion. Mark chordates (and tetrapods particularly) linguistically as the unusual case instead. with a little judicious editing, becomes:
Biologists typically talk of chordates, arthropods, molluscs, annelids, nematodes, rotifers, etc. individually; I still suspect there is some anthropocentric bias in our taxonomy (are we really different enough from chimps to justify a separate genus?), but far less than there is in the general population. Amusing example: At the Creation Museum, they have a chart of their taxonomy (as opposed to the standard biological taxonomy). It lists "man" as a separate category distinct from everything else, but then it ties together with a common ancestry groups as diverse as "dinosaurs", "insects", and (my personal favorite) "fungi". So the entire domain of fungi, from yeast to mushrooms, is allowed to have a common ancestor; but God forbid anyone suggest that humans and chimps are related.
I think there's a good argument that we are. (But I do not actually advocate a terminological change and am happy to let the biologists cut reality along whatever joints are convenient for them.) Edit: By "terminological change" I mean the one suggested in the linked quote.

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.


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 wh... (read more)

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 bein... (read more)

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.

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?

This is a more general problem. Why do some books get published, and others not? Why are some actors in Hollywood and others confined to shoestring-budget theater companies? As much as we like to think that capitalism is a meritocracy, in the real world merit often loses out to a lot of other factors, many of which we don't yet understand.

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.

I'm sure that's an important part of the answer. But still, how does the cascade get started? How does a book as singularly awful as Twilight become a million-seller?

Twilight is actually a great book, it's just that you are probably not its target audience. It may be misleading, because writing about vampires suggests that it's about mystery or fighting. Instead think about it as an erotic book for heterosexual women -- in this genre it is excellent. (And by pretending that it is not from the genre, it removes some stigma.)

The essence of the book is a description of an exceptionally attractive alpha male. Typical books from this genre are about princes or millionaires, but obviously vampires are stronger and richer than that, and also can be superhumanly attractive. If you read the book from this perspective, it actually makes a lot of sense (including the infinitely repeating descriptions of Edward's beautiful eyes, beautiful hands, beautiful feet, beautiful mouth, beautiful nose, etc.). And if you are the target audience, you don't need this explanation, you just feel the excitement, which you can later rationalize as good writing (and it helps that the writing actually is not too bad).

-- spoiler alert -- I agree, the explicit but g-rated eroticism was well done. However, to me the book series was mostly about the single unbelievably magical creature in it, Bella: * Despite being totally in love at 17, she manages to hold her own extremely well against a man who is much older, more experienced, smarter and more attractive. How often does that happen? * She is also selfless beyond all credibility (for example, having been unceremoniously dumped and left broken by the love of her life, she harbors no ill will toward him whatsoever and is ready to throw her life away to save the dude). Her first concern when she is in mortal danger is always about the people she loves, likes, barely knows, or doesn't know at all. Her own well-being is always an afterthought and she feels guilty that, by being a target, she endangers others. * Not a trace of an ego, either: she derives no satisfaction, only anguish from being pined for by two high-status men she likes. Every person I know would enjoy such a situation at least a little bit. (I'm ignoring the last book, where she did become a magical creature.)

Reminds me of Hanson - all that reads like serious signaling for being a good mother.

Edward is smarter? Oh, wait. Smarter than Bella. Low bar!


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 ... (read more)

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:

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'v... (read more)

1Eli Ace2y
The Psychological Foundations of Culture is my favorite. What are your other four?

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.


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... (read more)

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, ... (read more)

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 ... (read more)


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 Elie... (read more)

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 cont... (read more)

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.

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 ... (read more)

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 probl... (read more)

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 explan... (read more)

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.


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... (read more)

"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 abou... (read more)

Before deciding that Gould's theory was wrong or unimportant, read something from 2007.

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.

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 popul... (read more)

"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... (read more)

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... (read more)

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 ... (read more)

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 reorganize... (read more)

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 g... (read more)

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.

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... (read more)

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.


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... (read more)

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, ... (read more)

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. 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 differ

... (read more)
1David Bahry2y
Coyne had complicated feelings about Gould (but loved Gould's sometimes-collaborator Lewontin, who was his supervisor). Here's Coyne responding to Trivers' "Vignettes of famous evolutionary biologists, large and small":
1David Bahry2y
What mistake is that? I know there's a couple details that have been updated (e.g. the replicator / vehicle of selection distinction, from The Extended Phenotype, improved things) but overall he, like many other evolutionary biologists, still thinks it holds up really well

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... (read more)

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