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Thursday, March 17, 2005

Genuine controversies and the distracting nonsense of creationism

An interesting question and some complaints broke out in the comments last night: how can you have evolution without selection?

What's interesting about it is that this is coming from intelligent, informed supporters of good biology, and what it tells us is that our educational system is not doing its job. There are important concepts that should be taught in our public school system, and we're so caught up in this absolutely idiotic struggle with morons to keep stupid, disproven claims like "Evolution is false!" out of the curriculum that very few people are getting the fundamentals. Of course evolution proceeds in the absence of selection. There are even different kinds of selection.

All most people get in their education about evolution is the one, tiny idea of "survival of the fittest" and natural selection, and typically they only get a minute subset of that, the idea of directional selection. Populations interact with their environment and the weak are culled out, relentlessly driving the population towards some adaptive optimum. What's more, the most common mistake people with this idea make is to assume that removing the selective force means the population will stop evolving.

There is another form of selection, almost certainly more common than directional selection, called stabilizing selection. This process acts to eliminate forms that deviate from the norm, and tends to keep the properties of a population more constant. Remove this, by for instance putting the population under artificial conditions, like a lab, that allows or even encourages variants to flourish, and you get an increase in diversity.

And, of course, population genetics has all these fascinating mathematical formulas that reveal counterintuitive results, like that it is quite reasonable that even advantageous mutations to disappear from populations, or that deleterious alleles can become fixed. I could dig up some of this stuff for you, but I think pestering Reed Cartwright, who has done some nice work on his evomath series, could answer that even more effectively.

There are modes of change that are independent of selection. As a developmental biologist, I'm most interested in the intrinsic properties of developmental mechanisms that buffer organisms from change and also facilitate accommodation: variations in developmental genes don't necessarily mean the organism simply stops working or fails to develop, but instead, the flexibility of development means that either alternative processes stabilize the process, or more interestingly, the whole compensates to integrate the change in a continuous and non-disruptive manner. Brian Goodwin carries this idea to an extreme (to a point I find uncomfortable, actually, and I disagree with him on some things), but it's a similar idea that internal properties and physical factors dictate part of the features of an organism.

Stuart Kauffman is another source for the importance of physical and mathematical properties of the universe contributing to the parameters shaping life. Goodwin and Kauffman are out there on the bleeding edge, so I don't know that I'd necessarily endorse them as a place to start looking at the alternatives to selection…a better starting point would be SJ Gould. You don't have to read his monster Structure of Evolutionary Theory (although it would help: that book is all about the variety of modes of evolution), you can start small.

Try reading The Pleasures of Pluralism, an excellent critique of the excesses of the selectionists and in particular of evolutionary psychology, but also a paean to the other, too-often-neglected mechanisms of evolution.

In summary, Darwin cut to the heart of nature by insisting so forcefully that "natural selection has been the main, but not the exclusive means of modification"—and that hard-line adaptationism could only represent a simplistic caricature and distortion of his theory. We live in a world of enormous complexity in organic design and diversity—a world where some features of organisms evolved by an algorithmic form of natural selection, some by an equally algorithmic theory of unselected neutrality, some by the vagaries of history's contingency, and some as byproducts of other processes.

Another terrific and influential article by Gould and Lewontin is The Spandrels of San Marco. Again, they are deploring the narrow reductionism that blinkers much of molecular biology and evolutionary biology, but they are also promoting alternative ways of thinking.

An adaptationist programme has dominated evolutionary thought in England and the United States during the past 40 years. It is based on faith in the power of natural selection as an optimizing agent. It proceeds by breaking an organism into unitary 'traits' and proposing an adaptive story for each considered separately. Trade-offs among competing selective demands exert the only brake upon perfection; non-optimality is thereby rendered as a result of adaptation as well. We criticize this approach and attempt to reassert a competing notion (long popular in contental Europe) that organisms must be analyzed as integrated wholes, with Baupläne so constrained by phyletic heritage, pathways of development and general architecture that the constraints themselves become more interesting and more important in delimiting pathways of change than the selective force that may mediate change when it occurs. We fault the adaptationist programme for its failure to distinguish current utility from the reasons for origin (male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small); for its unwillingness to consider alternatives to adaptive stories; for its reliance upon plausibility alone as a criterion for accepting speculative tales; and for its failure to consider adequately such competing themes as random fixation of alleles, production of non-adaptive structures by developmental correlation with selected features (allometry, pleiotropy, material compensation, mechanically forced correlation), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. We support Darwin's own pluralistic approach to identifying the agents of evolutionary change.

(All emphases are mine)

Creationists lately have been babbling about "teaching the controversy". I'm all for that, but the controversy isn't what they think it is. None of these authors, not even the most radical, denies that evolution occurred. What they are arguing about is the relative importance of different modes of evolutionary change; and none of this discussion is filtering into our public schools, because a) teachers are intimidated from discussing evolution at all, and b) when they do, they have to address this ginned-up bogus bullshit promulgated by know-nothing creationists. If they were sincere about wanting to introduce controversial ideas into the classroom, they'd boot out everyone who even mentions that Intelligent Design creationism idiocy, and promote people who say they want to sink more hours into exploring the basic concepts of evolutionary biology more deeply. When do these bozos ever mention teaching about directional and stabilizing selection, or allometry, or pleiotropy, or developmental accommodation? Never. And that's the good stuff.


Gould SJ, Lewontin RC (1979) The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proc. R. Soc. Lond. B 205:581-598.


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Comments:
#18972: Danny Boy — 03/17  at  10:00 AM
Two opposable thumbs up! Thanks for sharing two wonderful essays by Gould, my favorite Darwinian popularizer. The spandrel paper, in particular, is quite a catch, since I've never read it before. Maybe you can also refer your readers to his close ally Niles Eldredge. In particular, I'd recommend two of his books, Reinventing Darwin and Patterns of Evolution. While not as grandiose in style, both are very informative and serve as a good counter-medicine to Dawkins and Dennett.



#18974: Michael Feldgarden — 03/17  at  10:12 AM
I think this is a really important point you made. To rephrase your point, there is a difference between the descent theory (i.e., evolution happens) and the theory of natural selection (one mechanism by which evolution can occur). I've always thought that if people knew about genetic drift (the random change and fixation of gene frequencies), that would be the serious theological challenge. In a way, evolution by natural selection can be very theologically/philosophically comforting: there is a certain rational logic to it.

But the idea that some variation becomes fixed (note to non-biologists: 'blue eye color has been "fixed" in the population' is 'biologist' for everyone has blue eyes) by random chance or if it's bad for you (even worse!), is not comforting. People want to know that life makes sense. The pseudo-mathematical arguments against evolution are strong evidence of this fear of randomness.



#18976: P.M.Bryant — 03/17  at  10:30 AM
Thanks for explaining this. I learned (or at least re-learned) quite a bit.



#18978: — 03/17  at  10:33 AM
A bit of clarification for Michael. The fact of evolution is not a "theory." It is an observed phenomena. The various theories of evolution seek to identify and explore the mechanisms that brings about the fact of evolution. For clarification, read this article by Gould:

http://www.stephenjaygould.org/library/gould_fact-and-theory.html



#18979: coturnix — 03/17  at  10:55 AM
Thank you for putting this clearly. I restrained myself from commenting last night. Lewontin is one of my heroes, too. Triple Helix, while not his best book, may be the best intro to his thought as it is a transcript of his three lectures that combine pretty much all of his thinking all in one place. Afterwards, one can explore his stuff in more detail in other books, or enjoy his "It Ain't Neccessarily So".

Blogwhoring alert:
http://sciencepolitics.blogspot.com/2004/12/wwdd-what-would-darwin-do-or_02.html
http://sciencepolitics.blogspot.com/2004/10/god-genes-and-conservatives.html
http://sciencepolitics.blogspot.com/2004/10/genocentrism-aids-anti-abortion.html



#18980: RPM — 03/17  at  10:57 AM
Good post. Though I doubt any IDiots or School Boards will pay attention. Motoo Kimura doesn't have a publicity group forcing elementary school kids to learn about the Neutral Theory.
And, of course, population genetics has all these fascinating mathematical formulas that reveal counterintuitive results, like that it is quite reasonable that even advantageous mutations to disappear from populations, or that deleterious alleles can become fixed.

If anyone is curious, the strength of natural selection depends on many factors: population size, environment, and other alleles in the population to name a few. Natural selection is not as effective in small populations as it is in large ones. Therefore, a deleterious allele can go to fixation in a small population due to neutral processes, even if the fitness of the allele predicts that it should be selected against and eventually lost.



#18981: — 03/17  at  10:57 AM
PZ, you can save your bandwidth. The AAAS has made the spandrel paper freely available here:

http://www.aaas.org/spp/dser/evolution/history/spandrel.shtml

Rrawr!



#18986: RPM — 03/17  at  11:56 AM
Coincidentally: I've got a Lewontin paper in front of me right now (on locating regions of evolutionary heterogeneity in DNA sequences). A bit too technical a read for most, but he's got a good statistical test that I'm thinking of applying to some data I've got. It's amazing that he can write for both a general audience and put out excellent technical papers.

And, as for the real debate. My advisor gave me some communications between Lewontin and Masatoshi Nei in which they discuss whether we can conclude alleles are under selection. These haven't been published -- they are copies of the actual letters that they sent to each other. Nei would bring up a neutral explanation for a particular example, and Lewontin would refute it with the evidence for selection (or evidence against neutrality). It really captures the essence of the "debate" amongst evolutionary geneticists. We all acknowledge that natural selection plays a role in evolution, but we can't agree on how important that role is.



#18999: — 03/17  at  01:26 PM
Thanks, PZ. I was always under the impression that natural selection is the only force involved.

Do you, uh, know of any papers on this that a high-schooler like me could understand?



#19003: — 03/17  at  02:05 PM
First, you are right on that biblical battles over nonsense interferes with improving education. Even us non-scientists who have read quite a bit of Gould still have to work to follow this type of information. One hurdle is the simple definition of the terminology. i.e. Using the lay meaning of selection (can be positive, negative, neutral, random, directed..... ) hinders my ability to easily see how evolution is not just a specific type of selection.

As I said before, good teachable moment; and now you have provided a nice path to learning more.

That learning more part is the best reason to hang out here.



#19004: — 03/17  at  02:11 PM
I'm pretty ignorant of biology, so it's likely someone else has mentioned my problem with the term "selection." This term implies an agency; that is, something selects. I understand that it is used to mean a particular process, but I think it might lead to misunderstanding of that process by laymen. It's fairly easy to understand how a genetic change that gives some advantage (in ability to attract a mate or find food or whatever) can lead to more offspring. But isn't it true that in this case "selection" means that the genetic change appears in offspring, and eventually, in a large part of the population, and then eventually in all of the population? If that is all there is to it, it seems easy to see how any random change, even one not particularly advantageous (or neutral, or even disadvantageous) might be passed to offspring. That "not particularly advantageous" change would then not be "selected" but would lead to an evolutionary change anyway. Or am I missing something?



#19005: — 03/17  at  02:44 PM
filteredvision - rather than papers, which by their nature tend to be quite narrow in focus, I would recommend a textbook on evolutionary biology. I have very high regard for Douglas Futuyma's textbook "Evolutionary Biology" which is now in its third edition. Hopefully you will be able to find it in a library near you. Its structure allows you to learn about different aspects of evolutionary theory without having to read the whole thing from cover to cover - although that wouldn't be a bad thing!



#19009: — 03/17  at  04:13 PM
I'm not a biologist by training, but I do not see how one can account for the apparent design we see in nature without acknowledging the force of natural selection.

I found Gould and Lewontin's infamous essay on spandrels more confusing than enlightening, but then I've never been one who cares much for explanation by way of metaphor.



#19011: — 03/17  at  04:31 PM
"[M]ale tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small."

Since Gould and Lewontin brought up the subject, I can't resist sharing what I think is the most interesting hypothesis for why those arms got so small.

It's like this: tyrannosaurs walked bipedally, with their trunk horizontal. Thus, their center of gravity had to be above their hind feet, and the weight of the "balanced beam" had to be equal on either side of the "fulcrum" -- the pelvis and tail on one side, and the chest, forelegs, and head on the other. Here's a skeletal reconstruction of T. rex, although it is shown lunging forward in an unbalanced way as if running fast. Now, this beast had a remarkably large and strong head, provided with powerful muscles. (Tooth marks on Triceratops and Edmontosaurus remains show that T. rex bit right through bones.) Therefore, its forequarters were heavy, even though its bones were hollow and its neck and chest were full of the sort of air sacs found in modern birds. The head grew heavier in successive tyrannosauroids -- compare Dilong paradoxus, an early member of the group, having a relatively smaller head and larger arms than T. rex. What's one way to reduce the weight of the front of the body, to keep balanced while walking? Reduce the size of the forelegs!

Thus, the hypothesis is that the tyrannosaur's forelegs grew smaller for weight compensation as the head grew larger. One way to test this hypothesis might be to compare whether the same effect ocurred in another lineage. Luckily, we have the carnosaurs -- see here for a cladogram showing their relationship to tyrannosauroids. They also grew larger over time, culminating in Giganotosaurus and Carcharodontosaurus, both of which may have been even larger than Tyrannosaurus. Compare this picture of Giganotosaurus (halfway down the page) to this one of the earlier carnosaur Allosaurus. The later species' head is somewhat larger in relation to its body, and its arms are somewhat smaller, though these changes have not reached nearly the extent they did in Tyrannosaurus.

Question for the biologists: if this hypothesis for the reduction of tyrannosaur arms is true, where does it fall in the list of mechanisms of evolution? Is it "mechanically forced correlation"?



#19015: — 03/17  at  05:39 PM
Thanks, PZ, very informative.



#19020: RPM — 03/17  at  06:24 PM
Male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small.

I believe that the current theory is that bipedalism is the ancestral state for dinosaurs. If that is the case, then the more interesting question would be, "How did brontosaurus get such big front legs?"



#19027: — 03/17  at  08:43 PM
I would think that as the prosauropods increased in gut size, the body developed for a more stable mode of moving around. Their arms were already developed enough for quadrupedal stances anyway, so they just "got pumped up!" *clap*.



#19028: Dan S. — 03/17  at  09:09 PM
A breath of fresh air after much time probably wasted arguing with an anti-evolutionist who depicted evolution as "the strong survive, the weak die, therefore if we believe evolution you get Nazis and infidelity.
-Dan S.



#19032: — 03/17  at  09:28 PM
Gould and Lewontin's paper is a rhetorical masterpiece but it too swings too far. For a very entertaining discussion of it take a look at:

Queller DC (1995) The spaniels of St. Marx and the Panglossian paradox: a
critique of the rhetorical. program. Quarterly Review of Biology 70: 485-89. [http://www.ruf.rice.edu/~evolve/pdf/Spaniels.pdf]



#19036: — 03/17  at  10:26 PM
RPM raises a point which prompted me to re-read Gould's essay "Of Kiwi Eggs and the Liberty Bell". The essay is all about the allometric changes that may occur in the relative sizes of different body parts as species in a lineage get bigger or smaller. I quote from it:
...if you ask me why kiwi eggs are so large, I reply, "Because kiwis are dwarfed descendants of larger birds, and just followed ordinary principles of scaling in their evolution." This answer differs sharply from the conventional form of evolutionary explanation: "Because these big eggs are good for something now, and natural selection favored them."

But are Tyrannosaurus forelimbs just the size you'd expect if you hugely scaled up a smallish bipedal ancestor similar to Dilong? Someone must have already done the calculations on this, but I don't know the reference. However, the comparison to Giganotosaurus (and other large theropods) suggests that Tyrannosaurus's forelimbs really are unusually diminutive. What's more, the ancestors of tyrannosauroids and carnosaurs had three fingers on their hands, Dilong had three fingers, and Giganotosaurus had three -- but Tyrannosaurus had only two. Something does seem to have favored reduction of the arms in this species, but my point was that it wasn't necessarily selection directly "for" small arms. You don't have to ask, "What was Tyrannosaurus doing with its arms that made two fingers better than three?" Instead, one possible solution is that small arms are simply a byproduct of a large head.



's avatar #19040: Nullifidian — 03/17  at  11:19 PM
For the sake of completeness, John Tooby and Leda Cosmides have a response to Gould's "The Pleasures of Pluralism."

"We are obliged, therefore, to spread the news, painful and bitter though it may be for some to hear, that all living things on earth are kindred.” Edward Abbey, Desert Solitaire



's avatar #19041: — 03/17  at  11:55 PM
Excellent article. Thanks. Evolution happens even in the absense of selection. Within the limits of the need of the organism to survive and reproduce. Everything is on the move and changing all the time, leading to a state of unending transformation. How simpler and nicer would all be if ID could be demonstrated true! But it is so obvious a nonsense.

Quod natura non sunt turpia



#19042: — 03/18  at  12:00 AM
You know, Prof. Myers, you are metamorphosing into one of the better essayists on evolution. Has Natural History called you yet?



's avatar #19043: Chris Clarke — 03/18  at  12:10 AM
You know, Prof. Myers, you are metamorphosing into one of the better essayists on evolution. Has Natural History called you yet?

You know, that's a pretty damned good idea.

"I do not think we should antagonize the religious when it is not warranted, though I think we should be willing to do so whenever it is.”
-- Glen Davidson



#19048: — 03/18  at  06:55 AM
This was an excellent post. I'm slightly partial to Stuart Kauffman myself and I thought the honorific "bleeding edge" was on the mark. In particular, in the last book I read, he presents a probablistic calculation/speculation as to how the mechanism shifting genetic instructions in cellular differentiation might work, which I dimly intuited. Does Prof. Myers have any remark on that account?



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