Darwinian fundamentalism, part 1

paulsamuel

Registered Senior Member
The New York Review of Books
June 12, 1997
Feature

Darwinian Fundamentalism, By Stephen Jay Gould

1.

With copious evidence ranging from Plato's haughtiness to Beethoven's tirades, we may conclude that the most brilliant people of history tend to be a prickly lot. But Charles Darwin must have been the most genial of geniuses. He was kind to a fault, even to the undeserving, and he never uttered a harsh wordÑor hardly ever, as his countryman Captain Corcoran once said. Darwin's disciple, George Romanes, expressed surprise at the only sharply critical Darwinian statement he had ever encountered: "In the whole range of Darwin's writings there cannot be found a passage so strongly worded as this: it presents the only note of bitterness in all the thousands of pages which he has published." Darwin directed this passage that Romanes found so striking against people who would simplify and caricature his theory as claiming that natural selection, and only natural selection, caused all evolutionary changes. He wrote in the last (1872) edition of The Origin of Species:

As my conclusions have lately been much misrepresented, and it has been stated that I attribute the modification of species exclusively to natural selection, I may be permitted to remark that in the first edition of this work, and subsequently, I placed in a most conspicuous positionÑnamely at the close of the IntroductionÑthe following words: "I am convinced that natural selection has been the main but not the exclusive means of modification." This has been of no avail. Great is the power of steady misrepresentation.

Darwin clearly loved his distinctive theory of natural selectionÑthe powerful idea that he often identified in letters as his dear "child." But, like any good parent, he understood limits and imposed discipline. He knew that the complex and comprehensive phenomena of evolution could not be fully rendered by any single cause, even one so ubiquitous and powerful as his own brainchild.

In this light, especially given history's tendency to recycle great issues, I am amused by an irony that has recently ensnared evolutionary theory. A movement of strict constructionism, a self-styled form of Darwinian fundamentalism, has risen to some prominence in a variety of fields, from the English biological heartland of John Maynard Smith to the uncompromising ideology (albeit in graceful prose) of his compatriot Richard Dawkins, to the equally narrow and more ponderous writing of the American philosopher Daniel Dennett (who entitled his latest book Darwin's Dangerous Idea).[1] Moreover, a larger group of strict constructionists are now engaged in an almost mordantly self-conscious effort to "revolutionize" the study of human behavior along a Darwinian straight and narrow under the name of "evolutionary psychology."

Some of these ideas have filtered into the general press, but the uniting theme of Darwinian fundamentalism has not been adequately stressed or identified. Professionals, on the other hand, are well aware of the connections. My colleague Niles Eldredge, for example, speaks of this coordinated movement as Ultra-Darwinism in his recent book, Reinventing Darwin.[2] Amid the variety of their subject matter, the ultra-Darwinists share a conviction that natural selection regulates everything of any importance in evolution, and that adaptation emerges as a universal result and ultimate test of selection's ubiquity.

The irony of this situation is twofold. First, as illustrated by the quotation above, Darwin himself strongly opposed the ultras of his own day. (In one sense, this nicety of history should not be relevant to modern concerns; maybe Darwin was overcautious, and modern ultras therefore out-Darwin Darwin for good reason. But since the modern ultras push their line with an almost theological fervor, and since the views of founding fathers do matter in religion, though supposedly not in science, Darwin's own fierce opposition does become a factor in judgment.) Second, the invigoration of modern evolutionary biology with exciting nonselectionist and nonadaptationist data from the three central disciplines of population genetics, developmental biology, and paleontology (see examples below) makes our pre-millennial decade an especially unpropitious time for Darwinian fundamentalismÑand seems only to reconfirm Darwin's own eminently sensible pluralism.
------------------------------------------------------------------------

Charles Darwin often remarked that his revolutionary work had two distinct aims: first, to demonstrate the fact of evolution (the genealogical connection of all organisms and a history of life regulated by "descent with modification"); second, to advance the theory of natural selection as the most important mechanism of evolution. Darwin triumphed in his first aim (American creationism of the Christian far right notwithstanding). Virtually all thinking people accept the factuality of evolution, and no conclusion in science enjoys better documentation. Darwin also succeeded substantially in his second aim. Natural selection, an immensely powerful idea with radical philosophical implications, is surely a major cause of evolution, as validated in theory and demonstrated by countless experiments. But is natural selection as ubiquitous and effectively exclusive as the ultras propose?

The radicalism of natural selection lies in its power to dethrone some of the deepest and most traditional comforts of Western thought, particularly the notion that nature's benevolence, order, and good design, with humans at a sensible summit of power and excellence, proves the existence of an omnipotent and benevolent creator who loves us most of all (the old-style theological version), or at least that nature has meaningful directions, and that humans fit into a sensible and predictable pattern regulating the totality (the modern and more secular version).

To these beliefs Darwinian natural selection presents the most contrary position imaginable. Only one causal force produces evolutionary change in Darwin's world: the unconscious struggle among individual organisms to promote their own personal reproductive successÑnothing else, and nothing higher (no force, for example, works explicitly for the good of species or the harmony of ecosystems). Richard Dawkins would narrow the focus of explanation even one step furtherÑto genes struggling for reproductive success within passive bodies (organisms) under the control of genesÑa hyper-Darwinian idea that I regard as a logically flawed and basically foolish caricature of Darwin's genuinely radical intent.

The very phenomena that traditional views cite as proof of benevolence and intentional orderÑthe good design of organisms and the harmony of ecosystemsÑarise by Darwin's process of natural selection only as side consequences of a singular causal principle of apparently opposite meaning: organisms struggling for themselves alone. (Good design becomes one pathway to reproductive success, while the harmony of ecosystems records a competitive balance among victors.) Darwin's system should be viewed as morally liberating, not cosmically depressing. The answers to moral questions cannot be found in nature's factuality in any case, so why not take the "cold bath" of recognizing nature as nonmoral, and not constructed to match our hopes? After all, life existed on earth for 3.5 billion years before we arrived; why should life's causal ways match our prescriptions for human meaning or decency?
------------------------------------------------------------------------

We now reach the technical and practical point that sets the ultra-Darwinian research agenda. Natural selection can be observed directly, but only in the unusual circumstances of controlled experiments in laboratories (on organisms with very short generations such as fruit flies) or within simplified and closely monitored systems in nature. Since evolution, in any substantial sense, takes so much time (more than the entire potential history of human observing!), we cannot, except in special circumstances, watch the process in action, and must therefore try to infer causes from resultsÑthe standard procedure in any historical science, by the way, and not a special impediment facing evolutionists.

The generally accepted result of natural selection is adaptationÑthe shaping of an organism's form, function, and behavior to achieve the Darwinian summum bonum of enhanced reproductive success. We must therefore study natural selection primarily from its resultsÑthat is, by concentrating on the putative adaptations of organisms. If we can interpret all relevant attributes of organisms as adaptations for reproductive success, then we may infer that natural selection has been the cause of evolutionary change. This strategy of researchÑthe so-called adaptationist programÑis the heart of Darwinian biology, and the fervent, singular credo of the ultras.

Since the ultras are fundamentalists at heart, and since fundamentalists generally try to stigmatize their opponents by depicting them as apostates from the one true way, may I state for the record that I (along with all other Darwinian pluralists) do not deny either the existence and central importance of adaptation, or the production of adaptation by natural selection. Yes, eyes are for seeing and feet are for moving. And, yes again, I know of no scientific mechanism other than natural selection with the proven power to build structures of such eminently workable design.

But does all the rest of evolutionÑall the phenomena of organic diversity, embryological architecture, and genetic structure, for exampleÑflow by simple extrapolation from selection's power to create the good design of organisms? Does the force that makes a functional eye also explain why the world houses more than five hundred thousand species of beetles and fewer than fifty species of priapulid worms? Or why most nucleotidesÑthe linked groups of molecules that build DNA and RNAÑin multicellular creatures do not code for any enzyme or protein involved in the construction of an organism? Or why ruling dinosaurs died and subordinate mammals survived to flourish and, along one oddly contingent pathway, to evolve a creature capable of building cities and understanding natural selection?

I do not deny that natural selection has helped us to explain phenomena at scales very distant from individual organisms, from the behavior of an ant colony to the survival of a redwood forest. But selection cannot suffice as a full explanation for many aspects of evolution; for other types and styles of causes become relevant, or even prevalent, in domains both far above and far below the traditional Darwinian locus of the organism. These other causes are not, as the ultras often claim, the product of thinly veiled attempts to smuggle purpose back into biology. These additional principles are as directionless, nonteleological, and materialistic as natural selection itselfÑbut they operate differently from Darwin's central mechanism. In other words, I agree with Darwin that natural selection is "not the exclusive means of modification."
------------------------------------------------------------------------

What an odd time to be a fundamentalist about adaptation and natural selectionÑwhen each major subdiscipline of evolutionary biology has been discovering other mechanisms as adjuncts to selection's centrality. Population genetics has worked out in theory, and validated in practice, an elegant, mathematical account of the large role that neutral, and therefore nonadaptive, changes play in the evolution of nucleotides, or individual units of DNA programs. Eyes may be adaptations, but most substitutions of one nucleotide for another within populations may not be adaptive.

In the most stunning evolutionary discoveries of our decade, developmental biologists have documented an astonishing "conservation," or close similarity, of basic pathways of development among phyla that have been evolving independently for at least 500 million years, and that seem so different in basic anatomy (insects and vertebrates, for example). The famous homeotic genes of fruit fliesÑresponsible for odd mutations that disturb the order of parts along the main body axis, placing legs, for example, where antennae or mouth parts should beÑare also present (and repeated four times on four separate chromosomes) in vertebrates, where they function in effectively the same way. The major developmental pathway for eyes is conserved and mediated by the same gene in squids, flies, and vertebrates, though the end products differ substantially (our single-lens eye vs. the multiple facets of insects). The same genes regulate the formation of top and bottom surfaces in insects and vertebrates, though with inverted orderÑas our back, with the spinal cord running above the gut, is anatomically equivalent to an insect's belly, where the main nerve cords run along the bottom surface, with the gut above.

One could argue, I suppose, that these instances of conservation only record adaptation, unchanged through all of life's vicissitudes because their optimality can't be improved. But most biologists feel that such stability acts primarily as a constraint upon the range and potentiality of adaptation, for if organisms of such different function and ecology must build bodies along the same basic pathways, then limitation of possibilities rather than adaptive honing to perfection becomes a dominant theme in evolution. At a minimum, in explaining evolutionary pathways through time, the constraints imposed by history rise to equal prominence with the immediate advantages of adaptation.

My own field of paleontology has strongly challenged the Darwinian premise that life's major transformations can be explained by adding up, through the immensity of geological time, the successive tiny changes produced generation after generation by natural selection. The extended stability of most species, and the branching off of new species in geological moments (however slow by the irrelevant scale of a human life)Ñthe pattern known as punctuated equilibriumÑrequires that long-term evolutionary trends be explained as the distinctive success of some species versus others, and not as a gradual accumulation of adaptations generated by organisms within a continuously evolving population. A trend may be set by high rates of branching in certain species within a larger group. But individual organisms do not branch; only populations doÑand the causes of a population's branching can rarely be reduced to the adaptive improvement of its individuals.
------------------------------------------------------------------------

The study of mass extinction has also disturbed the ultra-Darwinian consensus. We now know, at least for the terminal Cretaceous event some 65 million years ago that wiped out dinosaurs along with about 50 percent of marine invertebrate species, that some episodes of mass extinction are both truly catastrophic and set off by extraterrestrial impact. The death of some groups (like dinosaurs) in mass extinctions and the survival of others (like mammals), while surely not random, probably bears little relationship to the evolved, adaptive reasons for success of lineages in normal Darwinian times dominated by competition. Perhaps mammals survived (and humans ultimately evolved) because small creatures are more resistant to catastrophic extinction. And perhaps Cretaceous mammals were small primarily because they could not compete successfully in the larger size ranges of dominant dinosaurs. Immediate adaptation may bear no relationship to success over immensely long periods of geological change.

Why then should Darwinian fundamentalism be expressing itself so stridently when most evolutionary biologists have become more pluralistic in the light of these new discoveries and theories? I am no psychologist, but I suppose that the devotees of any superficially attractive cult must dig in when a general threat arises. "That old time religion; it's good enough for me." There is something immensely beguiling about strict adaptationismÑthe dream of an underpinning simplicity for an enormously complex and various world. If evolution were powered by a single force producing one kind of result, and if life's long and messy history could therefore be explained by extending small and orderly increments of adaptation through the immensity of geological time, then an explanatory simplicity might descend upon evolution's overt richness. Evolution then might become "algorithmic," a surefire logical procedure, as in Daniel Dennett's reverie. But what is wrong with messy richness, so long as we can construct an equally rich texture of satisfying explanation?
 
Darwinian Fundamentalism, parts 2 & 3

2.
Daniel Dennett's 1995 book, Darwin's Dangerous Idea, presents itself as the ultras' philosophical manifesto of pure adaptationism. Dennett explains the strict adaptationist view well enough, but he defends a miserly and blinkered picture of evolution in assuming that all important phenomena can be explained thereby. His limited and superficial book reads like a caricature of a caricatureÑfor if Richard Dawkins has trivialized Darwin's richness by adhering to the strictest form of adaptationist argument in a maximally reductionist mode, then Dennett, as Dawkins's publicist, manages to convert an already vitiated and improbable account into an even more simplistic and uncompromising doctrine. If history, as often noted, replays grandeurs as farces, and if T.H. Huxley truly acted as "Darwin's bulldog," then it is hard to resist thinking of Dennett, in this book, as "Dawkins's lapdog."

Dennett bases his argument on three images or metaphors, all sharing the common error of assuming that conventional natural selection, working in the adaptationist mode, can account for all evolution by extensionÑso that the entire history of life becomes one grand solution to problems in design. "Biology is engineering," Dennett tells us again and again. In a devastating review, published in the leading professional journal Evolution, and titled "Dennett's Dangerous Idea," H. Allen Orr notes:

His review of attempts by biologists to circumscribe the role of natural selection borders on a zealous defense of panselectionism. It is also absurdly unfairÉ. Dennett fundamentally misunderstands biologists' worries about adaptationism. Evolutionists are essentially unanimous thatÑwhere there is "intelligent Design"Ñit is caused by natural selectionÉ. Our problem is that, in many adaptive stories, the protagonist does not show dead-obvious signs of Design.

In his first metaphor, Dennett describes Darwin's dangerous idea of natural selection as a "universal acid"Ñto honor both its ubiquity and its power to corrode traditional Western beliefs. Speaking of adaptation, natural selection's main consequence, Dennett writes: "It plays a crucial role in the analysis of every biological event at every scale from the creation of the first self-replicating macromolecule on up." I certainly accept the acidic designationÑfor the power and influence of the idea of natural selection does lie in its radical philosophical contentÑbut few biologists would defend the blithe claim for ubiquity. If Dennett chooses to restrict his personal interest to the engineering side of biologyÑthe part that natural selection does constructÑthen he is welcome to do so. But he may not impose this limitation upon others, who know that the record of life contains many more evolutionary things than are dreamt of in Dennett's philosophy.

Natural selection does not explain why many evolutionary transitions from one nucleotide to another are neutral, and therefore nonadaptive. Natural selection does not explain why a meteor crashed into the earth 65 million years ago, setting in motion the extinction of half the world's species. As Orr points out, Dennett's disabling parochialism lies most clearly exposed in his failure to discuss the neutral theory of molecular evolution, or even to mention the name of its founder, the great Japanese geneticist Motoo KimuraÑfor few evolutionary biologists would deny that this theory ranks among the most interesting and powerful adjuncts to evolutionary explanation since Darwin's formulation of natural selection. You don't have to like the idea, but how can you possibly leave it out?

In a second metaphor, Dennett continually invokes an image of cranes and skyhooks. In his reductionist account of evolution, cranes build the good design of organisms upward from nature's physicochemical substrate. Cranes are good. Natural selection is evolution's basic crane; all other cranes (sexual reproduction, for example) act as mere auxiliaries to boost the speed or power of natural selection in constructing organisms of good design. Skyhooks, on the other hand, are spurious forms of special pleading that reach down from the numinous heavens and try to build organic complexity with ad hoc fallacies and speculations unlinked to other proven causes. Skyhooks, of course, are bad. Everything that isn't natural selection, or an aid to the operation of natural selection, is a skyhook.
------------------------------------------------------------------------

If you think that I am being simplistic or unfair to Dennett in this characterization, read his book and see if you can detect anything more substantial in this metaphor. I could only find a rhetorical stick for beating pluralists into line. Can't Dennett see that a third (and correct) option exists to his oddly dichotomous Hobson's choice: either accept the idea of one basic crane with auxiliaries, or believe in skyhooks. May I suggest that the platform of evolutionary explanation houses an assortment of basic cranes, all helping to build the edifice of life's history in its full grandeur (not only the architecture of well-engineered organisms). Natural selection may be the biggest crane with the largest set of auxiliaries, but Kimura's theory of neutralism is also a crane; so is punctuated equilibrium; so is the channelling of evolutionary change by developmental constraints. "In my father's house are many mansions"Ñand you need a lot of cranes to build something so splendid and variegated.

For his third metaphorÑthough he would demur and falsely label the claim as a fundamental statement about causesÑDennett describes evolution as an "algorithmic process." Algorithms are abstract rules of calculation, and fully general in making no reference to particular content. In Dennett's words: "An algorithm is a certain sort of formal process that can be counted onÑlogicallyÑto yield a certain sort of result whenever it is 'run' or instantiated." If evolution truly works by an algorithm, then all else in Dennett's simplistic system follows: we need only one kind of crane to supply the universal acid.

I am perfectly happy to allowÑindeed I do not see how anyone could denyÑthat natural selection, operating by its bare-bones mechanics, is algorithmic: variation proposes and selection disposes. So if natural selection builds all of evolution, without the interposition of auxiliary processes or intermediary complexities, then I suppose that evolution is algorithmic too. ButÑand here we encounter Dennett's disabling error once againÑevolution includes so much more than natural selection that it cannot be algorithmic in Dennett's simple calculational sense.

Yet Dennett yearns to subsume all the phenomenology of nature under the limited aegis of adaptation as an algorithmic result of natural selection. He writes: "Here, then, is Darwin's dangerous idea: the algorithmic level is the level that best accounts for the speed of the antelope, the wing of the eagle, the shape of the orchid, the diversity of species, and all the other occasions for wonder in the world of nature" (Dennett's italics). I will grant the antelope's run, the eagle's wing, and much of the orchid's shapeÑfor these are adaptations, produced by natural selection, and therefore legitimately in the algorithmic domain. But can Dennett really believe his own imperialistic extensions? Is the diversity of species no more than a calculational consequence of natural selection? Can anyone really believe, beyond the hype of rhetoric, that "all the other occasions for wonder in the world of nature" flow from adaptation?

Perhaps Dennett only gets excited when he can observe adaptive design, the legitimate algorithmic domain; but such an attitude surely represents a blinkered view of nature's potential interest. I regard the neutral substitution of nucleotides as an "occasion for wonder in the world of nature." And Imarvel at the probability that the impact of a meteor wiped out dinosaurs and gave mammals a chance. If this contingent event had not occurred, and imparted a distinctive pattern to the evolution of life, we would not be here to wonder about anything at all!

3.
"Straight is the gate, and narrow is the way." Fundamentalists of all stripes live by this venerable motto, and must therefore wield their unsleeping swords in constant mental fight against contrary opinions of apostates and opponents (who usually make up a sizable majorityÑfor, as Jesus also noted, "Wide is the gate, and broad is the way, that leadeth to destruction"). The favored fate for the nonelect varies, according to the temperament and power of true believers, from the kindness of simple pity to the refiner's fire of extirpation. But the basic ideological weapon of fundamentalism rarely departs much from the tried and true techniques of anathematization.

Unfortunately, at least for the ideals of intellectual discourse, anathematization rarely follows the dictates of logic or evidence, and nearly always scores distressingly high in heat/light ratio. Anathema also requires an anathemeeÑand I seem to have been elected. (Whatever my professional contributions to proper Darwinian pluralism, I stand convicted, Isuggest, primarily for my efforts to bring the full scope of technical debate, with all its complexities and messiness, but without loss of substance, to general readers.)

Personal attack generally deserves silence by way of response. But as two old troupers (Noam Chomsky and Salvador Luria) once advised me in my only comparable earlier incident, an exception must be made in one and only one circumstance: when denigrators float a demonstrably false charge that, if unanswered, may acquire a "life of its own." A false fact can be refuted, a false argument exposed; but how can one respond to a purely ad hominem attack? This harder, and altogether more discouraging, task may best be achieved by exposing internal inconsistency and unfairness of rhetoric.

John Maynard Smith, Emeritus Professor at Sussex and dean of British ultra-Darwinians, reviewed Dennett's book in this publicationÑthus providing small prospect for critical commentary. Maynard Smith began his supposed analysis of ultraDarwinian criticism with the following gratuitous remark:

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.[3]

It seems futile to reply to an attack so empty of content, and based only on comments by anonymous critics; if they were named, they would, Isuspect, turn out to be a very small circle of true believers. And if I may beg the editor's indulgence for one emotional outburst, may I say, at least, that I resent Maynard Smith's pompous offer of grudging acceptance for my utility in fighting creationism. I did not do so to win entry into his circle of genuine professionals (for I think that we both hold honored union cards therein), but rather as a member of the larger scientific community, and as a small contribution to the continual struggle of people who cherish rationality. We will not win this most important of all battles if we descend to the same tactics of backbiting and anathematization that characterize our true opponents.
------------------------------------------------------------------------

Instead of responding to Maynard Smith's attack against my integrity and scholarship, citing people unknown and with arguments unmentioned, let me, instead, merely remind him of the blatant inconsistency between his admirable past and lamentable present. Some sixteen years ago he wrote a highly critical but wonderfully supportive review of my early book of essays, The Panda's Thumb, stating: "I hope it will be obvious that my wish to argue with Gould is a compliment, not a criticism."[4] He then attended my series of Tanner Lectures at Cambridge in 1984 and wrote in a report for Nature, and under the remarkable title "Paleontology at the High Table," the kindest and most supportive critical commentary I have ever received. He argued that the work of a small group of American paleobiologists had brought the entire subject back to theoretical centrality within the evolutionary sciences.

The attitude of population geneticists to any palaeontologist rash enough to offer a contribution to evolutionary theory has been to tell him to go away and find another fossil, and not to bother the grownups.

In the last ten years, however, this situation has changed by the work of a group of palaeontologists, of whom Gould has been a leading figure.

He ended the article with a quintessential Oxbridge metaphor: "The Tanner lectures were an entertaining and stimulating occasion. The palaeontologists have too long been missing from the high table. Welcome back."

Maynard Smith then republished both papers (along with two others that cast my work in the central role of a section entitled "Did Darwin Get it Right?") in his 1988 volume of essays, Did Darwin Get it Right? Essays on Games, Sex and Evolution. Most remarkably of all, he then reviewed two books on dinosaurs for this journal and devoted more than half his space (much to the distress, I am sure, of the authors of the books supposedly under review) to a trenchant critique of my views on adaptation. He began by writing: "When, as often happens, I find myself dissenting from something written by Stephen Jay Gould, I remind myself that we share a common childhood experience. We were both dinosaur nuts." And he ended with an apology: "I fear that what started out as a review of two books about dinosaurs has wandered off into a discussion of the functional and adaptationist approaches to anatomy."[5]

So we face the enigma of a man who has written numerous articles, amounting to tens of thousands of words, about my workÑalways strongly and incisively critical, always richly informed (and always, I might add, enormously appreciated by me). But now Maynard Smith needs to canvass unnamed colleagues to find out that my ideas are "hardly worth bothering with." He really ought to be asking himself why he has been bothering about my work so intensely, and for so many years. Why this dramatic change? Has he been caught up in apocalyptic ultra-Darwinian fervor? I am, in any case, saddened that his once genuinely impressive critical abilities seem to have become submerged within the simplistic dogmatism epitomized by Darwin's Dangerous Idea, a dogmatism that threatens to compromise the true complexity, subtlety (and beauty) of evolutionary theory and the explanation of life's history. I shall examine this Darwinian fundamentalism further in a second, and concluding, article in the next issue.

Notes
[1] Darwin's Dangerous Idea: Evolution and the Meanings of Life (Simon and Schuster, 1995).
[2] Reinventing Darwin: The Great Debate at the High Table of Evolutionary Theory (John Wiley, 1995).
[3] The New York Review, November 30, 1995.
[4] London Review of Books, September 17-30, 1981.
[5] The New York Review, April 25, 1991.
 
Talk about counting angels dancing on pinheads. Still I agree that Dennet's book is patronising, narrow minded and riddled with assumptions.
 
about part 1.

I read part 1 now, and I must admit that I am quite disappointed with Gould. He accuses the Darwinian fundamentalist of stigmatising the opponent and Gould continues to do the same to them. So far his main points were that Darwin proposed that besides natural selection there were other mechanisms involved in evolution. Did anyone ever deny this? Did Darwin also not state that natural selection was the most important mechanism by far!

Darwin at his time also was dealing with a lot of uncertainty. He had no knowledge of genes or DNA and naturally he had to build in some uncertainty in his viewpoint. This can therefore hardly be used as an argument against the fundamentalist Darwinists.

What are Gould's other arguments so far.

Developmental biology: The conservation of important signalling-pathways throughout the animal kingdom. In what way this phenomenon would be unfavourable towards natural selection is unknown to me. Who ever claimed that evolution is not a natural history and that every generation has to work with what was present in the previous generation. How could any animal invent a new signaling pathway out of the blue for eye development if there already is one in place? As a developmental biologists I seriously think that this whole argument of development biology is misconstrued. He is trying to project the notion that the fundamentalist Darwinians are not seeing something that is obvious. But how can the fact that natural selection has to work with what is present be an argument against natural selection?

And then there is the chance argument. There are chance events that have a huge impact on the diversity of life, because some species groups didn't cope as well as others. Did anyone ever deny this? Is this the Holy Grail of evolution? A meteorite hitting a local environment killing of everything in a single location and causing the extinction of a single species because this is the only place they occur is a chance event. A huge impact killing of all dinosaurs' species worldwide, leaving absolutely none left, is a catastrophe. But is it a chance event? It is unfortunate for them, but other species managed to cope better with the changing environment. They were hit just as bad as the dinosaurs. They survived. Many species of their kind survived. Is this chance? Or is it because they happened to be better adapted to this radical environmental change?

In conclusion, so far (after just reading part 1) I think that gould is grasping at straws. His arguments are weak. His use of language aimed at hurting his opponents rather than building a real case.
Maybe part 2 will change my opinion. I will let you know.
 
part 2 and 3

Part2 and 3 is mostly about personal attacks and could be considered the equivalent of an extended and more pompous sciforum flamewar. I didn't really see anything constructive in these 2 parts. And so far I have been disappointed with Gould. The best he could come up with is that natural selection cannot explain why a meteor crashed into the earth 65 million years ago and causing a major extinction event, wiping out the dinosaurs. Is this his contribution then to the grand scheme of evolution? Is there all there is to it?

He didn't really address punctuated equilibrium here in detail, but I have the suspicion that this theory is just as shallow. Has anything interesting at all come out of all this questioning of natural selection? Natural selection still seems to be untouched if these are the best arguments. Or was Gould so selfish to take the penultimate argument to the grave?
 
Re: about part 1.

Originally posted by spuriousmonkey
So far his main points were that Darwin proposed that besides natural selection there were other mechanisms involved in evolution. Did anyone ever deny this?
Apparently, yes. Gould is saying that Dennet, Maynard-Smith and Dawkins are denying this.
Originally posted by spuriousmonkey
Did Darwin also not state that natural selection was the most important mechanism by far!
If we leave out the modality ("by far!") certainly Darwin thought this, and so does Gould.

Originally posted by spuriousmonkey
Darwin at his time also was dealing with a lot of uncertainty. He had no knowledge of genes or DNA and naturally he had to build in some uncertainty in his viewpoint. This can therefore hardly be used as an argument against the fundamentalist Darwinists.
Gould recognizes this and says so, but goes ahead anyway.

Originally posted by spuriousmonkey
Developmental biology: The conservation of important signalling-pathways throughout the animal kingdom. In what way this phenomenon would be unfavourable towards natural selection is unknown to me.
The point is that many structures, pathways, etc. are not due to adaptation through natural selection, but have arisen due to developmental constraints. This flys in the face of the adaptationists.

Originally posted by spuriousmonkey
But how can the fact that natural selection has to work with what is present be an argument against natural selection?
He is not arguing against natural selection. He is saying that much of the phenotypic diversity and evolution we see has arisen from mechanisms other than adaptation through natural selection.

Originally posted by spuriousmonkey
There are chance events that have a huge impact on the diversity of life, because some species groups didn't cope as well as others. Did anyone ever deny this?
Yes! This is completely anti-Darwinian. Natural selection works at the level of the organism, not group, according to whom Gould is calling ultra-Darwinians. This is Gould's point here, that selection can work at hierarchical levels, i.e. species selection, cladal selection, populational selection. This is entirely Gould's baby and he is nurturing it, much to the consternation of the ultra-Darwinians.

Originally posted by spuriousmonkey
A huge impact killing of all dinosaurs' species worldwide, leaving absolutely none left, is a catastrophe. But is it a chance event?
Of course a meteor impact is a chance event, evolutionarily-wise. There is no way for such short-lived organisms to form an evolutionary response to the periodicity of meteor impacts. The subsequent species sorting after the event is, in Gould's opinion, not chance, but due to species or cladal selection.
 
Re: part 2 and 3

Originally posted by spuriousmonkey
Is this his contribution then to the grand scheme of evolution? Is there all there is to it?
Ah,... no? Please see Gould's The Structure of Evolutionary Theory.

Originally posted by spuriousmonkey
He didn't really address punctuated equilibrium here in detail, but I have the suspicion that this theory is just as shallow. Has anything interesting at all come out of all this questioning of natural selection? Natural selection still seems to be untouched if these are the best arguments. Or was Gould so selfish to take the penultimate argument to the grave?
Goodness gracious! I didn't realize this essay would make you so angry. I thought it was thought provoking not the other kind of provoking. In regards to punctuated equilibrium, certainly a lot of research has been generated, don't know how interesting it is as that is a matter of personal opinion, but it's interesting to me.
 
Originally posted by paulsamuel
Ah,... no? Please see Gould's The Structure of Evolutionary Theory.

we are just talking about this essay, but if you want you can post the other main mechanisms that drive evolution here. I'm not going to read his books anymore. I'm fed up with his writing style.

i'm not angry, i'm just voicing my opinion on the essay. I'm just terribly disappointed that he makes so much fuss about nothing.
 
Last edited:
Re: Re: about part 1.

Originally posted by paulsamuel

The point is that many structures, pathways, etc. are not due to adaptation through natural selection, but have arisen due to developmental constraints. This flys in the face of the adaptationists.

arisen due to developmental constraints:

either this is a lot of hot air, or it is just what I said. Natural selection can only work with what is there, not with what would be optimal.

anyway...what do you think yourself about the essay. You just now repeated Goulds opinions in your replies. I also noticed that you just responded to some of my lines, instead of the thoughts. To be honest, I am not going to bother to reply to these remarks and just pick out some of your ideas. I hope you will do the same. After all, this is not a debate about who is right. This is a discussion.

group selection:

you mention that the extinction of entire groups of species is anti-darwinian because natural selection works on the level of the individual. That is very interesting of course, but these huge extinction events are caused by major changes in the environment (be it climate, new species etc). If the genetic information of a single individual cannot cope with the new environement at all, the chances are very high that another individual of the same species has similar problems. In fact, within a species all individual have rather similar genotypes and if there are major changes in the enviroment we can assume that all individual lack the same ability to cope with the new challenges. How this can possibly be anti-darwinian is beyond me. Let us be honest. These events can be explained on the level of the individual. It might not fit the fancy wording of a scholar of Goulds statue, but I am just a simple man and I like simple ideas.
 
Last edited:
Re: Re: Re: about part 1.

Originally posted by spuriousmonkey
arisen due to developmental constraints:

either this is a lot of hot air, or it is just what I said. Natural selection can only work with what is there, not with what would be optimal.
I don't understand your point here. Of course, "natural selection can only work with what is there," but just because it's there doesn't mean natural selection is working on it. There are characteristics of organisms that are not adaptive (not the result of natural selection).

Gould, in this essay, is right. In his words, " We live in a world of enormous complexity in organic design and diversityÑa 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. Why should such a complex and various world yield to one narrowly construed cause? "

Originally posted by spuriousmonkey
group selection:

you mention that the extinction of entire groups of species is anti-darwinian because natural selection works on the level of the individual. That is very interesting of course, but these huge extinction events are caused by major changes in the environment (be it climate, new species etc). If the genetic information of a single individual cannot cope with the new environement at all, the chances are very high that another individual of the same species has similar problems. In fact, within a species all individual have rather similar genotypes and if there are major changes in the enviroment we can assume that all individual lack the same ability to cope with the new challenges. How this can possibly be anti-darwinian is beyond me.
Because darwinian evolution is a gradual process of populational adaptive change through individual natural selection over geologic time spans. Evolution works at the population level, natural selection works at the individual level. Catastrophic extinction events are not darwinian evolution. This is fairly basic evolutionary theory.
 
what might be these features in organisms that are not adaptive. He didn't mention any in his essay.

Originally posted by paulsamuel

Because darwinian evolution is a gradual process of populational adaptive change through individual natural selection over geologic time spans. Evolution works at the population level, natural selection works at the individual level. Catastrophic extinction events are not darwinian evolution. This is fairly basic evolutionary theory.

The reference to geological timeframe is nonsensical. Natural selection doesn't have to work on geological timescales. That's just your opinion. It is clear that this isn't true. Speciation can occur in a single generation (references have been mentioned in some of the threads here). Selection pressure can be shortlived but substantial. This entire groupselection can be explained by natural selection as I have done in the previous post. If your only objection is the timescale then I am afraid that I do not find the postion of the group selectionist a very strong one.
 
Last edited:
Originally posted by spuriousmonkey
what might be these features in organisms that are not adaptive. He didn't mention any in his essay.
A couple of examples;

"The internal error of adaptationism arises from a failure to recognize that even the strictest operation of pure Darwinism builds organisms full of nonadaptive parts and behaviors. Nonadaptations arise for many reasons in Darwinian systems, but consider only my favorite principle of "spandrels."
All organisms evolve as complex and interconnected wholes, not as loose alliances of separate parts, each independently optimized by natural selection. Any adaptive change must also generate, in addition, a set of spandrels, or nonadaptive byproducts. These spandrels may later be "co-opted" for a secondary use. But we would make an egregious logical error if we argued that these secondary uses explain the existence of a spandrel. I may realize someday that my favorite boomerang fits beautifully into the arched space of my living room spandrel, but you would think me pretty silly if I argued that the spandrel exists to house the boomerang. Similarly, snails build their shells by winding a tube around an axis of coiling. This geometric process leaves an empty cylindrical space, called an umbilicus, along the axis. A few species of snails use the umbilicus as a brooding chamber for storing eggs. But the umbilicus arose as a nonadaptive spandrel, not as an adaptation for reproduction. The overwhelming majority of snails do not use their umbilicifor brooding, or for much of anything."

and,

"Direct adaptation is only one mode of evolutionary origin. After all, I also have nipples not because I need them, but because women do, and all humans share the same basic pathways of embryological development."

Originally posted by spuriousmonkey
The reference to geological timeframe is nonsensical. Natural selection doesn't have to work on geological timescales. That's just your opinion. It is clear that this isn't true. Speciation can occur in a single generation (references have been mentioned in some of the threads here). Selection pressure can be shortlived but substantial. This entire groupselection can be explained by natural selection as I have done in the previous post. If your only objection is the timescale then I am afraid that I do not find the postion of the group selectionist a very strong one.
It's not nonsensical. I was speaking of macroevoulutionary changes. I realize that microevolutionary processes can occur quickly, but this has nothing to do with what we're talking about. You did not explain group selection by natural selection in your previous post. If you think you did, you need to rephrase it and be more clear in your explanation. Start with how you're defining group selection, how natural selection at the individual level can lead to group selection, and how this supports the synthetic darwinian view of evolution (of which, I can assure you, they totally disagree with. The neo-darwinian synthesis, the proponents of which Gould is railing about, cannot support group selection, and if you think it does, you need to explain how).
 
Originally posted by paulsamuel

"Direct adaptation is only one mode of evolutionary origin. After all, I also have nipples not because I need them, but because women do, and all humans share the same basic pathways of embryological development."

I was pondering for a while on this and all I could think of was that this reflects to the basic 'which was first, the chicken or egg' problem. The mammalian nipple evolved as a Darwinian adaptation. Apparently it is too complicated or unnecessary to remove the nipple in the man. The nipple is therefore a classic example of a mammalian adaptiation. If you concentrate on the example of male nipples then a paradox occurs. It is seemingly not a Darwinian adaptation. But it is of course, the nipple has arisen as a Darwinian adaptation. Indeed males have them because it is apparently impossible to separate the developmental pathways. But is this really an example of anti-Darwinian evolution? I have to admit that I am not convinced at all.

Similarly for
Originally posted by paulsamuel

Similarly, snails build their shells by winding a tube around an axis of coiling. This geometric process leaves an empty cylindrical space, called an umbilicus, along the axis. A few species of snails use the umbilicus as a brooding chamber for storing eggs. But the umbilicus arose as a nonadaptive spandrel, not as an adaptation for reproduction. The overwhelming majority of snails do not use their umbilicifor brooding, or for much of anything."
The space might not have a function, but it is generated during the process of creating a shell. How is having a protective shell not adaptive?
Also for many structures we are just not aware of the function, although they might have one.

Originally posted by paulsamuel

It's not nonsensical. I was speaking of macroevoulutionary changes. I realize that microevolutionary processes can occur quickly, but this has nothing to do with what we're talking about. You did not explain group selection by natural selection in your previous post. If you think you did, you need to rephrase it and be more clear in your explanation. Start with how you're defining group selection, how natural selection at the individual level can lead to group selection, and how this supports the synthetic darwinian view of evolution (of which, I can assure you, they totally disagree with. The neo-darwinian synthesis, the proponents of which Gould is railing about, cannot support group selection, and if you think it does, you need to explain how).

Group selection: I don't really believe in group selection if there would be specific mechanisms required other than natural selection for this kind of selection. But if you ask me how selection on the level of a group can apparently occur, then I would basically have to repeat my earlier explanation. Within a species there is genetic variation. Between two different species the genetic variation is however on a completely different level. In fact, the variation between species is usually much greater than the variation within a species. The level of variation between a group of related species is usually less than the level of variation between unrelated species, or groups of unrelated species. A specific group of species will usually share some specific characteristics or adaptations based on shared genetic information.
During rapid changes in environment selection still works on the individual level, but because a group of similar species shares roughly similar genetic information, they will have similar difficulties to adapt to changing environments. An entire group of species can have a disadvantage based on some specific shared genetic information or a broad set of genetic information. The question is if this is a matter of chance. Of course a species is unlucky if none of the individuals has the capacity to adapt. And of course a species cannot prepare itself for the unknown future. But this unfortunate situation is not limited to catastrophic events only. The same is the case for periods of 'normal' evolution.
Evolution during catastrophic events could therefore just be seen as periods in which regular evolution is speeded up. Chance is a factor of evolution at all times. But the chances that a species will survive the new environment are much higher if it possess the genetic variation to deal with the new environment. Natural selection to the extreme.

I don't know if this supports the synthetic darwinian view, since I only really care about my own view on evolution. I'm not going to defend someone else his view. I can only defend mine. I personally I don't really see how natural selection cannot explain all events that we have discussed. There might be other mechanisms, but I think that so far Gould has only mentioned examples that show why in his opinion natural selection cannot explain everything. I would like to hear something more constructive though, before I switch camps. What is the mechanism for group selection? And what is then the mechanism behind non-Darwinian adaptations.
 
Spookz - you're not allowed to suggest this. Species are supposed to survive and evolve without actually caring whether they live or die, and certainly not by intentionally competing with each other. The idea is unscientific.
 
it is a shame to see this thread slide away to the bottom of the page. And in reflection I have to admit that I didn't really know enough about group selection to make any of the bold statements I have made. I came to this conclusion after glancing over some of the terms used in the field of group selection. Apparently my views are probably quite amateurish. Not that this ever stopped me before of course.
 
God Gametes

I discussed the arguments put forward by Steven Jay Gould in my ebook “God Gametes and the Planet of the Butterfly Queen.” The God Gametes concept is definitely a sky hook but is not supportive of any theological interpretations of the meaning of life. Cranes can not build a universe or the gravitational forces that hold it all together. So if there is a sky hook out there that done that then my bet is that it also had a hand in kick-starting life on earth and evolving it into more and more complex forms. My ebook can be downloaded free from www.e-publishingaustralia.com but the section on Gould is posted below.

Post 1.

Stephen Jay Gould

We now look at how Steven Jay Gould addresses the issue of probability in Life’s Grandeur. Gould is a noted Darwinist who also dismisses the suggestion that life was designed. He is strongly opposed to any theory supportive of a creator but does not agree with the gradualist approach of Richard Dawkins. Gould does not try to explain the existence of matter in our universe or the origin of gravitational forces regulating it. He suggests that life:

“presumably began in primeval oceans as a result of sequential chemical reactions based on original constituents of atmospheres and oceans, and regulated by principles of physics for self-organizing systems.” 9

But again we are left wondering about the origin of the ‘primeval oceans’ and how ‘atmosphere’ might have formed in which life got started. It is also a little difficult to know what caused these ‘sequential chemical reactions’ or to comprehend a force that can formulate ‘principles of physics’ and drive the ‘self organization’ of matter.
Gould rejects a notion of progress in evolution or any arguments that our species has a ‘preferred status’ or is the ‘culmination’ of some creative process. But by rejecting a creator or a purpose in life, he needs to present a model for explaining the organised complexity so abundant on our planet.


The Left Wall:

Gould uses an analogy when outlining his case, asking us to imagine a drunk walking down the street. He depicts the drunk as the progress of life in that there is no purpose or direction to his actions or which way he will fall. We are asked to imagine that there is a wall on his left and a gutter to his right. The left wall is ‘minimum complexity’ and the gutter, ‘improved design’. He then points out that there is only one way the drunk can fall. He will most likely wobble to the left as many times as to the right but when he hits the left wall he will bounce back. If he keeps going he will eventually fall and it will be into the gutter. Gould then argues that there is such a brick wall to life on earth, his rationale being that there is a minimum level of complexity necessary for life to exist. He seems to suggest that when creatures hit this wall they bounce back and take the only option available and this is to evolve greater complexity.10
He assumes there is a level of complexity supporting life yet the Darwinian paradigm does not provide any mechanism for supporting minimum complexity. It is true that if the complexity of creatures falls below a certain level they will cease to live. But ceasing to live is different from bouncing back. What is it that stops the level of complexity falling below the minimum required to support life? Gould cannot say that it is the property of life, or by analogy a left brick wall, for Darwinists argue that there is no preferred status for life to exist.
Individuals can die and species can become extinct so a more accurate Darwinian analogy would be a ‘left cliff’. Any creature that devolved below the minimum level of complexity needed to support life would fall over this cliff never to be seen again.
We know that 99.9% of all life that ever lived on earth has become extinct.11 This would suggest that there is no left wall designed to prevent living creatures from facing the perils of extinction. It might also be worthwhile looking at why so many species have suffered this fate. The left wall analogy implies that these are species that fell to the left and did not bounce back, yet this does not appear to be the case. Most unsuccessful species are those that fell the other way. They attempted to make a living by evolving greater complexity and failed to adapt when environmental conditions changed.
Gould’s analogy has not explained why species tend to evolve greater complexity when there is no left wall bouncing them in that direction. He should also acknowledge that falling into the gutter is not a good analogy for evolving complexity. A better comparison would be to suggest that our drunk suddenly did a triple pirouette off the back foot, leaped 30 feet into the air and landed amidst oncoming traffic on other side of the street. This would paint a clearer picture of the developmental processes involved and also suggest a reason for the high incidents of extinctions associated with evolving greater complexity.

Continued in God Gametes Post 2.
 
Last edited:
Back
Top