Thursday, July 03, 2008

Good Science Writers: Richard Lewontin

 
Richard "Dick" Lewontin (1929 - ) is Alexander Agassiz Research Professor at Harvard University (Boston, USA). He is a well-known geneticist and the discoverer of extensive variation in organisms at the molecular level, with John Hubby [Citation Classic]. (See The Cause of Variation in a Population.)

Lewontin is also one of the authors of the textbook Modern Genetic Analysis. It's impossible to tell who wrote what in that textbook but I strongly suspect that Lewontin is responsible for the material on Random Genetic Drift and Population Size that I quoted last year.

He is the co-author (with Stephen Jay Gould) of The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist program, one of the most important papers in evolutionary biology [Citation Classic]. If you haven't carefully read that paper then you should do so right now.

In addition to his genetics textbook, Lewontin has also written books on The genetic basis of evolutionary change (1974), Human diversity (1995) and The Triple Helix: Gene, Organism, and Environment (2000). He is probably best known for his crusade against genetic determinism as described in Not in Our Genes: Biology, Ideology and Human Nature (with Steven Rose and Leon Kamin) as well as other books. Like several of his colleagues, Lewontin is a frequent contributer to the New York Review of Books and some of his best work has been republished in It Ain't Necessarily So: The Dream of the Human Genome and Other Illusions (2000).

Richard Dawkins did not choose anything from Richard Lewontin for The Oxford Book of Modern Science Writing. I can't imagine why.

The first selection is from Biology as Ideology: The Doctrine of DNA (1991) based on a series of lectures he gave at the University of Toronto in 1990. I was there.

The subject is sociobiology and genetic determinism. He has just finished explaining why there's no such thing as universal human nature—at least not the sort that is postulated by evolutionary psychologists and sociobiologists.
The final step in the sociobiological argument is to say that the genes we possess for universal human nature have been established in us through evolution by natural selection. That is, once upon a time human beings varied genetically in the degree to which they were aggressive, xenophobic, indoctrinable, male dominant, and so on, but those individuals who were most aggressive or most male dominant left more offspring, so the genes that were eventually left in us as a species were the ones that now determine those traits. The argument of natural selection seems a fairly simple and straightforward one for some kinds of traits. For example, it is argued, the more aggressive of our ancestors would leave more offspring because they would swoop down on the less aggressive and eliminate them. The more entrepreneurial would have appropriated more resources in short supply and starved out the wimps. In each of these cases, it is easy to make up a plausible story that would explain the superior reproductive abilities of one type over another.

There are, however, some traits that are said to be universal and that do not lend themselves so easily to this story of individual reproductive advantage. An example, and one that is discussed a great deal by sociobiologists, is altruistic behavior. Why should we be cooperative under some circumstances, and why should we sometimes give up what appear to be immediate advantage for the benefit of others? To explain altruism, sociobiologists advance the theory of kin selection. Natural selection for a trait does not require that individuals possessing it leave more offspring but only that the genes coding the trait be represented in larger numbers in future generations.

There are two ways to increase the representation of one's genes in future generations. One is to leave more offspring. The other is to arrange that even if one does not leave more offspring, one's relative's do so, since close relatives share genes. So, a person could sacrifice his reproduction completely, provided his brothers and sisters left many more children. Thus, his kind of genes would increase indirectly through his relatives and, in this indirect way, he would leave more offspring. An example of this phenomenon is the occurrence of "helpers at the nest" in birds, in which it is said that nonreproductive birds help out their close relatives, who are then able to raise more than the ordinary number of offspring and in the end more family genes are left. To make kin selection work, a sufficient number of excess offspring must be left by relatives. For example, if an individual gives up its own reproduction, its brothers and sisters must have twice as many offspring as ordinarily, but one can at least tell a story that might make this plausible.

We are then left with those traits that do not even benefit relatives differentially, for example, a general altruism toward all members of the species. Why are we good to strangers? For this phenomenon, the sociobiologist provides the theory of "reciprocal altruism." The argument is that even if we are unrelated, if I do you a favor that costs me something, you will remember that favor and reciprocate in the future, and by this indirect path I will succeed in advancing my own reproduction. An example often given is that of a drowning person. You see someone drowning and jump in to save that person even at the risk of your own life. In the future, when you are drowning, the person whose life you have saved will remember, and save you in gratitude. By this indirect path you will increase your own probability of survival and reproduction over the long run. The problem with this story is, of course, that the last person you would want to depend on to save you when you are drowning is someone whom you had to save in the past, since he or she is not likely to be a strong swimmer.

The real difficulty with the process of explanation that allows direct advantage, or kin selection, or reciprocal altruism when one or the other is useful in the explanation, is that a story can be invented that will explain the natural selective advantage of any trait imaginable. When we combine individual selective advantage with the possibility of kin selection and reciprocal altruism, it is hard to imagine any human trait for which a plausible scenario for its selective advantage could not be invented. The real problem is to find out whether any of these stories is true. One must distinguish between plausible stories, things that might be true, and true stories, things that actually have happened. How do we know that human altruism arose because of kin selection or reciprocal altruistic selection? At the very minimum, we might ask whether there is any evidence that such selective processes are going on at the present, but in fact no one has ever measured in any human population the actual reproductive advantage or disadvantage of any human behavior. All of the sociobiological explanations of the evolution of human behavior are like Rudyard Kipling's Just So stories of how the camel got his hump and how the elephant got its trunk. They are just stories. Science has turned into a game.
The second example is from a review of Darwinism Defended: A Guide to the Evolution Controversies by Michael Ruse. It was first published in The New York Review of Books on June 16, 1983 and reprinted in It Ain't Necessarily So: The Dream of the Human Genome and Other Illusions.
If Darwin's revolution was not in proclaiming evolution as a fact, then it must have been in his theory of its mechanism. And what was that theory? Why, "natural selection," of course, which then makes the theory of natural selection the very essence of Darwinism and any doubt about the universal efficacy of natural selection anti-Darwinian. There is a form of vulgar Darwinism, characteristic of the late nineteenth century and rejuvenated in the last ten years, which sees all aspects of shape, function, and behavior of all organisms as having been molded in exquisite detail by natural selection—the greater survival and reproduction of those organisms whose traits make them "adapted" for the struggle for existence. This Panglossian view is held largely by functional anatomists and comparative physiologists who, after all, make a living by explaining what everything is good for, and by sociologists who are self-consciously trying to win immortality by making their own small revolution. Evolutionary geneticists, on the other hand, who have spent the last sixty years in detailed experimental and theoretical analysis of the actual process of evolutionary change, and most epistemologists take a more pluralistic view of the forces driving evolution.

An occasional philosopher has allied himself or herself with the "adaptationists," who give exclusive emphasis to natural selection., and one such, Michael Ruse, makes a characteristic presentation in Darwinism Defended. Darwinism, the representative of objective modern science, is under ideologically motivated attack. Professor Ruse is alarmed: "'Darwinism,' as I shall refer to Darwin-inspired evolutionary thought, is threatened from almost every quarter." Well, not from every quarter, just the right and left flanks, it seems. First, the fundamentalists, supported by Ronald Regan, make a know-nothing assault from the right. No sooner have real evolutionists wheeled to face this attack than they are fallen upon by subversive elements from the left, "biologists with Marxist sympathies" and their "fellow travelers" among philosophers who argue "that any evolutionary theory based on Darwinian principles—particularly any theory that sees natural selection as the key to evolutionary change—is misleadingly incomplete."

Onto the field, mounted upon his charger perfectly adapted for the purpose, with weapons carefully shaped by selection to spread maximum confusion among the enemy, not to mention innocent civilians, comes Professor Ruse, "trying to rescue ... from the morass into which so many seem determined to drag them," "Darwin's life and achievements." In all fairness to Professor Ruse, he did not invent this version of events. The theory that evolutionary science is being brutally beaten and cut with crosses, hammers, and sickles made its first appearance in E.O. Wilson's On Human Nature as the only plausible explanation he could imagine for the failure of sociobiology to achieve instant, universal, and lasting adherence. The situation of evolutionary theory, however, is rather more complex and more interesting than Professor Ruse's Manichaean analysis suggests....

What vulgar Darwinists fail to understand, however, is that there is an asymmetry in Darwin's scheme. When adaptation is observed, it can be explained by the differential survival and reproduction of variant types being guided and biased by their differential efficiency or resistance to environmental stresses and dangers. But any cause of differential survival and reproduction, even when it has nothing to do with the struggle for existence, will result in some evolution, just not adaptive evolution.

The Panglossians have confused Darwin's discovery that all adaptation is a consequence of variational evolution with the claim that all variation evolution leads to adaptation. Even if biologists cannot, philosophers are supposed to distinguish between the assertion that "all x is y" and the assertion that "all y is x," and most have. This is not simply a logical question but an empirical one. What evolutionary geneticists and developmental biologists have been doing for the last sixty years is to accumulate a knowledge of a variety of forces that cause the frequency of variant types to change, and that do not fall under the rubric of adaptation by natural selection. These include, to name a few: random fixation of nonadaptive or even maladaptive traits because of limitations of population size and the colonization of new areas by small numbers of founders; the acquisition of traits because the genes influencing them are dragged along on the same chromosome as some totally unrelated gene that is being selected; and developmental side effects of genes that have been selected for some quite different reason.
.

[Photo credit: The photograph of Richard Lewontin is from (Photographs of Participants in the Molecular Evolution Workshop)]

28 comments:

  1. This comment has been removed by the author.

    ReplyDelete
  2. Lewontin is fantastic. You should blog about Isadore Nabi, vs. Dawkins et al.

    ReplyDelete
  3. These last two posts on Good Science Writers make fascinating reading.

    ReplyDelete
  4. Lewontin is pretty good as both a scientist and a popularizer of science. But sections of Not In Our Genes - namely, the Marxoid parts - are pathetic, more befitting a hemp fiber-attired granola punk with a Che Guevara poster on his dorm wall than distinguished scholars.

    And I very much doubt that Lewontin would approve of Dr. Moran's views on alleged cognitive differences between human races.

    Tupaia

    ReplyDelete
  5. My supervisor did his post-doc in Dr. Lewontin's lab in the 70s and speaks very highly of him as both a scientist and a writer. I TA a class taught by my boss in biology called "Human Diversity and Human Nature", which employs a lot of writing by Gould, Lewontin, and others in order to examine the reality of genetic bases for human racial and intelligence classification (and thus prejudices).

    Although I do enjoy most of Lewontin's writing, I must agree with Tupiaa in that Not in Our Genes is not one of his finest works. There is a serious undercurrent of political socialism in it that has nothing to do with rational scientific analysis. The book also vastly misrepresents The Selfish Gene (Dawkins discusses this in the notes to the 1989 and 30th anniversary editions of his book), and I'm not entirely surprised that Dawkins left him out of his new book...

    ReplyDelete
  6. The Gould and Lewontin Spandrels paper is what got me interested in evolutionary biology.

    ReplyDelete
  7. Tupaia says,

    And I very much doubt that Lewontin would approve of Dr. Moran's views on alleged cognitive differences between human races.

    What exactly, are you referring to? I doubt very much that Lewontin and I would disagree.

    ReplyDelete
  8. Larry Moran: "What exactly, are you referring to?"

    If I understand you correctly based on past posts and comments about Watson, Stent etc. you believe that races are real biological categories, that IQ difference between people is partly genetic, and that races (must, due to drift and selection) differ in frequencies of these genes. Therefore, races must differ to some extent in genetic potential for intelligence.

    Am I mistaken?

    http://sandwalk.blogspot.com/2007/10/is-there-genetic-component-to.html

    http://sandwalk.blogspot.com/2008/06/gunther-stent-1924-2008.html

    Lewontin, in contrast, believes that races are invalid biological categories and is dubious about the role of genes in IQ differences between races (and classes, nations etc.). (Am on correct about this?)

    Anyone else want to join in on Dr. Moran's and Dr. Lewontin's respective views on the matter? Sanders? Greg Laden?

    Tupaia

    ReplyDelete
  9. http://sandwalk.blogspot.com/2008/01/matt-nisbet-asks-embarrassing-question.html

    Tupaia: "It is an empirical fact that races have different mean IQs. Do human races have different genetic potential for intelligence?"

    Dr. Moran: "I don't know whether different races have different average IQ's. It's not unreasonable that they could since there is clearly a genetic component to intelligence and there is variation between individuals that appears to due to different alleles.

    It seems extremely unlikely that the frequencies of these allele would be exactly the same in all races."

    ----------------------

    Note:

    Actually, I shouldn't have stated it "races have different mean IQs." I should have wrote "in some studies races have different mean IQ."

    In fact, IQ differences between races change, disappear, or are reversed, depending on the study. I believe that patterns of racial IQ differences reflect socioeconomic, cultural, and other environmental differences, which themselves can have recurrent patterns, resulting in the appearance of "racial" differences.

    Tupaia

    ReplyDelete
  10. The Spandrels paper is one of my all-time favorites. Lewontin also made one of my favorite quips. Considering his views on genes and behavior, a postdoc of his once asked him what he taught in his Genes-and-Behavior course at Harvard. "Basic behavior genetics of simple organisms", Lewontin replied, then added,"But I certainly didn't say that mutants in Drosophila are why I hate lima beans".

    (The story is related in Jonathan Weiner's book "Time, Love, Memory, p. 218)

    ReplyDelete
  11. The reason that Dawkins didn't put a Lewontin selection in his anthology is pretty obvious. The two men don't like each other at all. Lewontin, for instance, published a scathing (and misguided) review of The Selfish Gene when it first appeared. Ever since then Dawkins has had no use for the man.

    ReplyDelete
  12. Re: Tupaia

    You claim that Moran's views on IQ/race is: "IQ difference between people is partly genetic, and that races (must, due to drift and selection) differ in frequencies of these genes."

    But, the quote you provide to demonstrate his views do *not* show that he believes there is a genetic IQ difference between the races. Based on the quotes, it seems that Moran is *agnostic* about genetic IQ differences between the races. He neither claims there are no differences between the races, nor claims that there are differences.

    ReplyDelete
  13. Lewontin: "... no one has ever measured in any human population the actual reproductive advantage or disadvantage of any human behavior."

    Unfortunately, whether the above is true or not (it is trivially easy to show it is false), Lewontin did his best to try to prevent inquiry into these types of questions, calling it false and dangerous science, and he was not above smearing the reputation of people who tried to research the question (just ask E.O. Wilson). [See "Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond" by Ullica Segerstrale.] It's one thing to say that one "side" of an issue is getting it wrong, it is another (and very shameful) thing to say that merely investigating something is wrong.

    And IMO, anyone who argues against strawman hypotheses (as he does above in his characterization of the reciprocal altruism using a drowning scenario) is not being an honest writer.

    And Tupaia is correct about the difference in Lewontin and Moran's views about intelligence, as at least Larry considers it a scientifically legitimate question (regardless of what view Larry actually has about the answer).

    ReplyDelete
  14. Bayesian Bouffant, FCDMonday, July 07, 2008 10:44:00 AM

    From I Wish I'd made You Angry Earlier by Max F. Perutz; Cold Spring Harbor Press, expanded edition 2003:

    (p. 251:) The Harvard University geneticist Richard Lewontin's book (It Ain't Necessarily So)consists of review essays originally published in the New York Review of Books between 1981 and 1993, with epilogues to bring them up to date or to answer his critics. They range from developmental biology to intelligence tests, human sexual behaviour, cloning and the human genome. They are erudite, readable, and they debunk a great deal of hype. The are also full of factual errors, prejudiced misjudgements and vital omissions...

    (p. 254:) Lewontin's black sheep include molecular biology, the human genome and gene therapy. He attacks Max Delbruck, the physicist who pioneered the immensely fruitful genetics of bacterial viruses, and whom he wrongly describes as a pupil of Schrodinger. He brands the phage group, the enthusiastic band of young people whom Delbruck assembled around himself, as "a political apparatus," and molecular biology as "a religion," which is absurd. He is right when he ridicules molecular biologist Walter Gilbert's vision of the human genome as its "holy grail" that will change our philosophic understanding of ourselves, but he then continues: "It is a sure sign of their alienation from revealed religion that a scientific community with a high concentration of Eastern European Jews and atheists has chosen for its central metaphor the most mystery-laden object of medieval Christianity." Remarks about people's race, religion and origin have no place in a book about science.

    (p. 257:) The man who invented the chemical method used for sequencing the genome was Fred Sanger, not Allan Maxam and Walter Gilbert whom Lewontin credits with it...

    (Maxam and Gilbert sequencing is based on specific chemical cleavage of nucleotides. Sanger sequencing is based on termination of template extension with dideoxy analogs. The technology used for the human genome project does indeed derive from Sanger's method.)

    ReplyDelete
  15. "Richard Dawkins did not choose anything from Richard Lewontin for The Oxford Book of Modern Science Writing. I can't imagine why."

    I love it when darwinists "fight".

    ReplyDelete
  16. This comment has been removed by the author.

    ReplyDelete

  17. Lewontin: "... no one has ever measured in any human population the actual reproductive advantage or disadvantage of any human behavior."

    Unfortunately, whether the above is true or not (it is trivially easy to show it is false),


    It is perfectly all right to pose as of yet untested hypotheses of course. (As it is also all right to skew the assumptions based on some likelihood for different mechanisms. Research programs must start somewhere, or at times concentrate on good methods.)

    But it isn't evidence for a factual science area. As I'm a layman I'm curious if there is any such evidence for sociobiology or evolutionary psychology as of yet.


    the acquisition of traits because the genes influencing them are dragged along on the same chromosome as some totally unrelated gene that is being selected;


    Another layman question, as I don't know the debate.

    This part starts out as a seeming criticism of natural selection as a mechanism with "universal efficacy". Then it ends with chalking up selective sweeps as no evidence for "adaptation by natural selection".

    Is this a sleight of hand, or is Lewontin really just attacking adaptionists instead of "selectionists", if you excuse my naive terminology? (If the later, he is really, really so underhanded by describing the simple meaning of implication just previously...)

    @ Divalent:


    And IMO, anyone who argues against strawman hypotheses (as he does above in his characterization of the reciprocal altruism using a drowning scenario) is not being an honest writer.


    I reacted too, but mainly because I hadn't figured on the down side on not allowing selection for a particular "trait" (swimming?) as a problem for altruism. After all, it is highly unlikely that a swimmer would face that particular problem later, but instead need help with his "bad traits".

    So I don't feel like Lewontin is making the strawman under pretensions of disregarding it. The next paragraph describes "the real difficulty". But it looks like an indirect smearing under the guise of making an inappropriate argument to accentuate a valid point. [Phew!]

    And anyway it is cut, that part doesn't look like "good science writing" to me.

    [Eek! Dunno how people can speculate on such specious traits anyway. I feel like I need a ... swim.]

    ReplyDelete
  18. Lewontin gets close to the truth on many accounts but does not acknowledge the centrality of having a theory of organism. Without this, placing the focus on population genetics (his field) will always be misleading: the organism is collapsed between the gene level and the population level.

    In toher words to gte a correct view of evolution Lewontin would have to renounce to population gentics as the basic framework and thus be willing to erase the blacknoard for a new beggining point

    (Notice that by erasing the blacboard I do not mena to destroy the achievements of population genetic and the questions it is capable of answering, but merely, not to consider it as starting point of all evolutionary understanding)

    ReplyDelete
  19. Lewontin: "... no one has ever measured in any human population the actual reproductive advantage or disadvantage of any human behavior."

    (Divalent:) Unfortunately, whether the above is true or not (it is trivially easy to show it is false)

    (Torbjörn:) It is perfectly all right to pose as of yet untested hypotheses of course. [...] But it isn't evidence for a factual science area. As I'm a layman I'm curious if there is any such evidence for sociobiology or evolutionary psychology as of yet.

    Those are fields of study. Do you mean evidence for specific hypotheses within those fields? Lewontin's claim is much more sweeping though. As for why he's trivially wrong: celibacy is a human behaviour, and it's trivially easy to show a reproductive disadvantage from celibacy (for example, measure reproductive output of nuns compared to a control group)

    As for less obvious examples, pre-industrial Finnish fishing communities had a higher survival rate of twins; a human behaviour (fishing) provided a reproductive advantage.

    It is much more difficult to pinpoint particular human behaviours as adaptations (in the above examples, fishing and not being celibate aren't adaptations), but "proving" an adaptation is difficult in any species.

    ReplyDelete
  20. of course, what would be interesting to the adaptationist is a human behavior that is shown to be genetically variable (selectable). This for sure is what Lewontin means.

    ReplyDelete
  21. Larry:
    Lewontin would fart in your face if he heard your stupid endorsments of Stent and Watson's bullshit. And that's that!

    ReplyDelete
  22. of course, what would be interesting to the adaptationist is a human behavior that is shown to be genetically variable (selectable). This for sure is what Lewontin means.

    As a great science writer he should have been able to easily express that, instead of the completely different claim he made now.

    As for your suggestion, variation at the DRD4 locus appears to be associated with human personality traits, like novelty seeking.

    ReplyDelete
  23. How about a meta-analysis of all data, Windy?


    Biol Psychiatry. 2008 Jan 15;63(2):197-206. Epub 2007 Jun 15.Click here to read Links
    Association of the dopamine D4 receptor (DRD4) gene and approach-related personality traits: meta-analysis and new data.
    Munafò MR, Yalcin B, Willis-Owen SA, Flint J.

    Department of Experimental Psychology, University of Bristol, Bristol, United Kingdom. marcus.munafo@bristol.ac.uk

    BACKGROUND: Two variants in the dopamine D4 receptor (DRD4) gene have been reported to be associated with human approach-related traits such as novelty seeking and extraversion. However, the strength of evidence for this association remains uncertain. METHODS: We conducted a meta-analysis of published studies of the association between the DRD4 gene variable number of tandem repeats (VNTR) and C-521T polymorphisms and human approach-related personality traits, including novelty seeking, extraversion, and impulsivity, restricted to adult samples recruited from nonpsychiatric populations, and extended on this literature by attempting to confirm any evidence of association in a replication sample (n = 309) selected for extreme scores on the extraversion subscale of the Eysenck Personality Questionnaire from a large (n = 40,090) population-based sample. RESULTS: Our initial meta-analysis supported the association of the DRD4 C-521T polymorphism, but not the VNTR polymorphism, with approach-related traits. This conclusion was qualified by evidence of significant publication bias and the failure to detect association in a replication sample comprising individuals at the extremes of the trait distribution. The association of the C-521T polymorphism observed in our initial meta-analysis was robust to the inclusion of these new data, but our revised meta-analysis indicated that the association was present for measures of novelty seeking and impulsivity but not for measures of extraversion. CONCLUSIONS: The DRD4 gene may be associated with measures of novelty seeking and impulsivity but not extraversion. The association of the C-521T variant with these measures, if genuine, may account for up to 3% of phenotypic variance.



    3%, Windy. FUCKIN 3 PERCENT

    In other words: The usual horsepooey.

    Of course if you had a little more antireductionist theoretical insight, you could realize beforehand, that assigning much influence of a single gene to a trait like "novelty seeking" is, ehem, unlikely.

    ReplyDelete
  24. Aren't you always asking why, if gradualism is true, people don't find any mutations of small effect? And now that there is one, you think it's "horsepooey"? Fucking moron.

    Of course if you had a little more antireductionist theoretical insight, you could realize beforehand, that assigning much influence of a single gene to a trait like "novelty seeking" is, ehem, unlikely.

    Moron. I didn't say "much influence", I said "appears to be associated with". This study supports my statement despite your attempt at spin.

    ReplyDelete
  25. hahaha. Sore loser?
    I guess with you darwin-maniacs, the smaller the effect, the better, huh?

    No use for u, windy. Byebye

    ReplyDelete
  26. I honestly feel sorry for u, windy, so, I'll stoop to explain stupidity.

    "Aren't you always asking why, if gradualism is true, people don't find any mutations of small effect?"

    You are an idiot. Of course genes with small effects exist (although, many times, like this case, the effect is so small the evidence is only provisional) I have never denied this.

    Whta we hardly ever find is evidence that a given trait is an accumulation by selection of sevral such genes.

    Notice what an asshole you are: If the amount had been, say 50%, you'd be throwing a party: gentic detemrination of personality!! if I had shown you 0.0001%...party too! it just proves darwinian gradualism.

    ReplyDelete
  27. the truth is, the smaller the effect of a gene, the more unlikely it is that it will have any affect on fitness (positive or negative), and not so, "the better for darwinism" as poor windy here has fooled himself into believing.

    ReplyDelete
  28. Life intervened, but finally returning to old threads FWIW:

    Thanks, windy!

    ReplyDelete