These themes recur throughout this section, and they
introduce
Ecology of Genes (6.
Richard-Dawkins-The-Devil-s-Chaplain
In hard-fought examples we may notice a progressive shift in resources from other parts of the animal's economy
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to service the arms race. And in any case the improvement in equip-
ment will normally be progressive. Another kind of positive feedback in evolution, if R. A. Fisher and his followers are right, results from sexual selection. Once again, progressive evolution is the expected consequence.
Progressive increase in morphological complexity is to be expected only in taxa whose way of life benefits from morphological complexity. Progressive increase in brain size is to be expected only in animals where braininess is an advantage. This may, for all I know, constitute a minority of lineages. But what I do insist is that in a majority of evolutionary lineages there will be progressive evolution towards some- thing. It won't, however, be the same thing in different lineages (this was the point about swifts and elephants). And there is no general reason to expect a majority of lineages to progress in the directions pioneered by our human line.
But have I now defined progress so generally as to make it a blandly useless word? I don't think so. To say that the evolution of the verte- brate eye was progressive is to say something quite strong and quite important. If you could lay out all the intermediate ancestors in chronological order you'd find that, first, for a majority of dimensions
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of measurement, the changes would be transitive over the whole sequence. That is, if A is ancestral to B which is ancestral to C, the direction of change from A to B is likely to be the same as the direction of change from B to C. Second, the number of successive steps over which progress is seen is likely to be large: the transitive series extends beyond A, B and C, far down the alphabet. Third, an engineer would judge the performance to have improved over the sequence. Fourth, the number of separate features combining and conspiring to improve performance would increase. Finally, this kind of progress really matters because it is the key to answering the Hoyle challenge. There will be exceptional reversals, for instance in the evolution of blind cave fish, where eyes degenerate because they are not used and are costly to make. And there will doubtless be periods of stasis where there is no evolution at all, progressive or otherwise.
To conclude this point, Gould is wrong to say that the appearance of progress in evolution is a statistical illusion. It does not result just from a change in variance as a baseball-style artefact. To be sure, complexity, braininess and other particular qualities dear to the human ego should not necessarily be expected to increase progressively in a majority of lineages - though it would be interesting if they did: the investigations
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of McShea, Jerison and others are not a waste of time. But if you
define progress less chauvinistically - if you let the animals bring their own definition - you will find progress, in a genuinely interesting sense of the word, nearly everywhere.
Now it is important to stress that, on this adaptationist view (unlike the 'evolution of evolvability' view to be discussed shortly), progressive evolution is to be expected only on the short to medium term. Coevolutionary arms races may last for millions of years, but probably not hundreds of millions. Over the very long timescale, asteroids and other catastrophes bring evolution to a dead stop, major taxa and entire radiations go extinct. Ecological vacuums are created, to befilledby new adaptive radiations driven by new ranges of arms races. The several arms races between carnivorous dinosaurs and their prey were later mirrored by a succession of analogous arms races between carnivorous mammals and their prey. Each of these successive and separate arms races powered sequences of evolution which were progressive in my sense. But there was no global progress over the hundreds of millions of years, only a sawtooth succession of small progresses terminated by extinctions. Nonetheless, the ramp phase of each sawtooth was properly and significantly progressive.
Ironically for such an eloquent foe of progress, Gould flirts with the idea that evolution itself changes over the long haul, but he puts it in a topsy-turvy way which has undoubtedly been widely misleading. It is
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more fully expounded in Wonderful Life but reprised in the present book. For Gould, evolution in the Cambrian was a different kind of process from evolution today. The Cambrian was a period of evolution- ary 'experiment', evolutionary 'trial and error', evolutionary 'false starts'. It was a period of 'explosive' invention, before evolution stabilized into the humdrum process we see today. It was the fertile time when all the great 'fundamental body plans' were invented. Nowadays, evolution just tinkers with old body plans. Back in the Cambrian, new phyla and new classes arose. Nowadays we only get new species!
This may be a slight caricature of Gould's own considered position,
but there is no doubt that the many American nonspecialists who
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unfortunately, as Maynard Smith wickedly observes, get their
evolutionary knowledge almost entirely from Gould, have been deeply misled. Admittedly, what follows is an extreme example, but Daniel Dennett has recounted a conversation with a philosopher colleague who read Wonderful Life as arguing that the Cambrian phyla did not have a common ancestor - that they had sprung up as independently initiated life forms! When Dennett assured him that this was not Gould's claim, his colleague's response was, 'Well then, what is all the fuss about? '
Even some professional evolutionists have been inspired by Gould's rhetoric into committing some pretty remarkable solecisms. Leakey and
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Lewin's The Sixth Extinction is an excellent book except for its Chapter
3, 'The Mainspring of Evolution', which is avowedly heavily influenced by Gould. The following quotations from that chapter could hardly be more embarrassingly explicit:
Why haven't new animal body plans continued to crawl out of the evolutionary cauldron during the past hundreds of millions of years?
In early Cambrian times, innovations at the phylum level survived because they faced little competition.
Below the level of the family, the Cambrian explosion produced relatively few species, whereas in the post-Permian a tremendous species diversity burgeoned. Above family level however, the post-Permian radiation faltered, with few new classes and no new phyla being generated. Evidently, the mainspring of evolution operated in both periods, but it propelled greater extreme experimentation in the Cambrian than in the post-Permian, and greater variations on existing themes in the post-Permian.
Hence, evolution in Cambrian organisms could take bigger leaps, including phylum-level leaps, while later on it would be more constrained, making only modest jumps, up to the class level.
It is as though a gardener looked at an old oak tree and remarked,
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wonderingly: 'Isn't it strange that no major new boughs have appeared on this tree recently. These days, all the new growth appears to be at the twig level! '
As it happens, molecular clock evidence indicates that the 'Cambrian Explosion' may never have happened. Far from the major phyla diverging from a point at the beginning of the Cambrian, Wray,
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Levinton and Shapiro present evidence that the common ancestors of
the major phyla are staggered through hundreds of millions of years back in the Precambrian. But never mind that. That is not the point I want to make. Even if there really was a Cambrian explosion such that all the major phyla diverged during a ten million-year period, this is no reason to think that Cambrian evolution was a qualitatively special kind of super-jumpy process. Bauplane don't drop out of a clear Platonic sky, they evolve step by step from predecessors, and they do so (I bet, and so would Gould if explicitly challenged) under approximately the same Darwinian rules as we see today.
'Phylum-level leaps' and 'modest jumps, up to the class level' are the sheerest nonsense. Jumps above the species level don't happen, and nobody who thinks about it for two minutes claims that they do. Even the great phyla, when they originally bifurcated one from another, were just pairs of new species, members of the same genus. Classes are species that diverged a very long time ago, and phyla are species that diverged an even longer time ago. Indeed it is a moot - and rather empty - question precisely when in the course of the step-by-step, gradual mutual divergence of, say, mollusc ancestors and annelid ancestors after the time when they were congeneric species, we should wish to say that the divergence had reached 'Bauplan' status. A good case could be made that The Bauplan is a myth, probably as pernicious as any of the myths that Stephen Gould has so ably combatted, but this one, in its modern form, is largely perpetuated by him.
I return, finally, to the 'evolution of evolvability' and a very real sense in which evolution itself may evolve, progressively, over a longer timescale than the individual ramps of the arms race sawtooth. Notwith- standing Gould's just scepticism over the tendency to label each era by its newest arrivals, there really is a good possibility that major innovations in embryological technique open up new vistas of evolutionary possibility and that these constitute genuinely progressive improvements. * The
*This is the idea that I dubbed 'The Evolution of Evolvability' (in C. Langton (ed. ), Artificial Life (Santa Fe, Addison Wesley, 1982)) and Maynard Smith and Szathmary wrote a book about 0. Maynard Smith and E. Szathmary, The Major Transitions in Evolution (Oxford, W. H. Freeman/Spektrum, 1995)).
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origin of the chromosome, of the bounded cell, of organized meiosis, diploidy and sex, of the eucaryotic cell, of multicellularity, of gastrulation, of molluscan torsion, of segmentation - each of these may have constituted a watershed event in the history of life. Not just in the normal Darwinian sense of assisting individuals to survive and reproduce, but watershed in the sense of boosting evolution itself in ways that seem entitled to the label progressive. It may well be that after, say, the invention of multicellularity, or the invention of segmentation, evolution was never the same again. In this sense there may be a one-way ratchet of progressive innovation in evolution.
For this reason over the long term, and because of the cumulative character of coevolutionary arms races over the shorter term, Gould's attempt to reduce all progress to a trivial, baseball-style artefact con- stitutes a surprising impoverishment, an uncharacteristic slight, an unwonted demeaning of the richness of evolutionary processes.
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Unfinished Correspondence with a Darwinian Heavyweight
The following correspondence was never completed and now, sadly, it never can be.
9 December 2001
Stephen Jay Gould Harvard
Dear Steve
Recently I received an email from Phillip Johnson, founder of the so-called 'Intelligent Design' school of creationists, crowing in triumph because one of his colleagues, Jonathan Wells, had been invited to take part in a debate at Harvard. He included the text of his email on his 'Wedge of Truth' web site, in which he announced the Wells debate under the headline 'Wells Hits a Home Run at Harvard'.
http://www. arn. org/docs/pjweekly/pj_weekly_011202. htm
The 'Home Run' turns out to be NOT a resounding success by Wells in convincing the audience, NOR any kind of besting of his opponent (Stephen Palumbi, who tells me he agreed to take part, with great reluctance, only because somebody at Harvard had ALREADY invited Wells and it was too late to do anything about that). There is no suggestion that Wells did well in the debate, nor even any obvious interest in whether he did. No, the 'Home Run' was simply and solely the matter of being invited by Harvard in the first place. These people have no hope of convincing reputable scientists by their ridiculous arguments. Instead, what they seek is the oxygen of respectability. We give them this oxygen by the mere act of ENGAGING with them at all. They don't mind being beaten in argument. What matters is that we give them recognition by bothering to argue with them in public.
You convinced me of this years ago when I phoned you up (you have probably
forgotten this) to ask your advice when I was invited to debate Duane P Gish. 218
? Ever since that phone call, I have repeatedly cited you and refused to debate these people, not because I am afraid of 'losing' the debate, but because, as you said, just to appear on a platform with them is to lend them the respectability they crave. Whatever might be the outcome of the debate, the mere fact that it is staged at all suggests to ignorant bystanders that there must be something worth debating, on something like equal terms.
First, I am interested to know whether you still hold to this view, as I do. Second, I am proposing that you might consider uniting with me (no need to involve others) in signing a short letter, say to the New York Review of Books, explaining publicly why we do not debate creationists (including the 'Intelligent Design' euphemism for creationists) and encouraging other evolutionary biologists to follow suit.
Such a letter would have great impact precisely because there have been widely publicised differences, and even animosities, between us (differences which creationists, with extreme intellectual dishonesty, have not hesitated to exploit). And I would not suggest writing a long disquisition on the technical differences which remain between us. That would only confuse the issue, make it harder to agree on a final draft, and lessen the impact. I wouldn't even mention our differences. I suggest a brief letter to the editor, explaining why we do not engage with 'intelligent design' or any other species of creationists, and offering our letter as a model for others to cite in refusing such invitations in the future. We both have better things to do with our time than give it over to such nonsense. Having just reached my sixtieth birthday (we are almost exactly the same age) I feel this keenly.
Steve replied on 11th December 2001, a warm and friendly email enthusiastically agreeing that a joint letter was an excellent idea, and saying that he'd be delighted to join me as the sole other signatory. He agreed that the New York Review of Books might well be the best place, and proposed that I should write a first draft. I reproduce that here, exactly as I sent it to him for his approval.
14 December 2001
Dear Editor
Like any flourishing science, the study of evolution has its internal controversies, as we both know. But no qualified scientist doubts that evolution is a fact, in the ordinarily accepted sense in which it is a fact that the Earth orbits the Sun. It is a fact that human beings are cousins
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to monkeys, kangaroos, jellyfish and bacteria. No reputable biologist doubts this. Nor do reputable theologians, from the Pope on. Unfortunately, many lay Americans do, including some frighteningly influential, powerful and, above all, well-financed ones.
We are continually invited to engage in public debates against creationists, including latter-day creationists disguised under the euphemism 'Intelligent Design Theorists'. We always refuse, for one overriding reason. If we may be allowed to spell this reason out publicly, we hope our letter may be helpful to other evolutionary scientists plagued by similar invitations.
The question of who would 'win' such a debate is not at issue. Winning is not what these people realistically aspire to. The coup they seek is simply the recognition of being allowed to share a platform with a real scientist in thefirstplace. This will suggest to innocent bystanders that there must be material here that is genuinely worth debating, on something like equal terms.
At the moment of writing, the leading 'Intelligent Design' website
reports a debate at Harvard under the banner headline 'Wells Hits a
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Home Run at Harvard'. Jonathan Wells is a creationist, incidentally a
long-time devotee of the Unification Church (the 'Moonies'). * He had a debate last month against Stephen Palumbi, Professor of Biology at Harvard University. 'Home Run' might seem to suggest that Reverends (sic) Wells scored some kind of victory over Professor Palumbi. Or at least that he made powerful points and his speech was well received. No such claim is made. It doesn't even seem to be of interest.
The 'Home Run' turns out to be simply the public demonstration at Harvard that, in the words of the website's author, Phillip Johnson, 'This is the sort of debate that is now occurring in universities. ' There was a victory, but it occurred long before the debate itself. The creationist scored his home run at the moment the invitation from Harvard landed on his doormat. It came, by the way, not from any biological, or indeed scientific, department, but from the Institute of Politics.
Phillip Johnson himself, founding father of the 'Intelligent Design'
''Darwinism: Why I went for a Second Ph. D. ' is Jonathan Wells' own testimony on the turning point of his life: 'Father's words, my studies, and my prayers convinced me that I should devote my life to destroying Darwinism, just as many of my fellow Unificationists had already devoted their lives to destroying Marxism. When Father chose me (along with about a dozen other seminary graduates) to enter a Ph. D. program in 1978, I welcomed the opportunity to prepare myself for battle. ' ('Father', of course, is the Moonies' name for Rev Moon himself). http://www. tparents. org/LibraryAJnincation/Talks/Wells/DARWIN. htm.
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? movement (not a biologist, nor a scientist of any kind, but a lawyer who became a mid-life born-again Christian), wrote, in a letter of 6 April 2001, which he copied to one of us:
It isn't worth my while to debate every ambitious Darwinist who wants to try his hand at ridiculing the opposition, so my general policy is that Darwinists have to put a significant figure at risk before I will agree to a debate. That means specifically Dawkins or Gould, or someone of like stature and public visibility.
Well, we can condescend too, and we have the advantage that evolutionary scientists don't need the publicity such debates can bring. In the unlikely event that a significant argument should ever emerge from the ranks of creationism/'intelligent design', we will be happy to debate it. Meanwhile, we shall cultivate our evolutionary gardens, occasionally engaging in the more exacting and worthwhile task of debating each other. What we shall not do is abet creationists in their disreputable quest for free publicity and unearned academic respectability.
In all humility, we offer these thoughts to our colleagues who receive similar invitations to debate.
Unfortunately, Steve never got around to revising the letter, which therefore lacks the stylish panache which his dexterous touch would have lent it. I received one further email, apologizing for the delay and hoping to deal with the matter soon. The subsequent silence, I now realize, coincided with his last illness. I therefore offer my draft, imperfect as it is, in the hope that it may go some way towards conveying the message which I originally learned from him many years ago. It is my sincere hope that he would have approved the content of the letter, but of course I cannot be sure.
To close this section on a note of such concord may seem puzzling.
Given that Steve was as much a neo-Darwinist as I am, what did we dis-
agree about? The major disagreement emerges clearly out of his last big
5
book, The Structure of Evolutionary Theory,TM which I had no opportunity
to see until after his death. It is appropriate, therefore, to spell out that issue here, and it also, as it happens, forms a natural bridge to the next essay. The question under dispute is this: what is the role of genes in evolution? Is it, to use Gould's phrase, 'book-keeping or causation'?
Gould saw natural selection as operating on many levels in the hierarchy of life. Indeed it may, after a fashion, but I believe that such selection can have evolutionary consequences only when the entities selected consist of 'replicators'. A replicator is a unit of coded information, of high fidelity but occasionally mutable, with some causal power over its own fate. Genes are
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such entities. So, in principle, are memes, but they are not under discus- sion here. Biological natural selection, at whatever level we may see it, results in evolutionary effects only insofar as it gives rise to changes in gene frequencies in gene pools. Gould, however, saw genes only as 'book- keepers', passively tracking the changes going on at other levels. In my view, whatever else genes are, they must be more than book-keepers, otherwise natural selection cannot work. If a genetic change has no causal influence on bodies, or at least on something that natural selection can 'see', natural selection cannot favour or disfavour it. No evolutionary change will result.
Gould and I would agree that genes can be seen as a book in which is
written the evolutionary history of a species. In Unweaving the Rainbow I
called it 'The Genetic Book of the Dead'. But the book is written via the
natural selection of randomly varying genes, chosen by virtue of their
causal influence on bodies. Book-keeping is precisely the wrong metaphor,
because it reverses the causal arrow, almost in Lamarckian fashion, and
makes the genes passive recorders. I dealt with this in 1982 (The Extended
Phenotype) in my distinction between 'active replicators' and 'passive
replicators'. The point is also explained in David Barash's superb review of
136 Gould's book.
Book-keeping is perversely - and characteristically - a valuable meta- phor precisely because it is so diametrically back to front. Not for the first time, the characteristic vividness and clarity of a Gouldian metaphor helps us to see vividly and clearly what is wrong with the Gouldian message - and how it needs to be reversed in order to get at the truth.
I hope this brief note will not be seen as taking advantage to get the last word. The Structure of Evolutionary Theory is such a massively powerful last word, it will keep us all busy replying to it for years. What a brilliant way
for a scholar to go. I shall miss him.
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THERE IS ALL AFRICA AND HER PRODIGIES IN US
I am one of those (it includes most people who have ever spent time south of the Sahara) who think of Africa as a place of enchantment. For me it stems from faint but haunting childhood memories, coupled with the mature understanding that Africa is our ancestral home.
These themes recur throughout this section, and they introduce Ecology of Genes (6. 1), my Foreword to Harvey Croze and John Reader's Pyramids of Life. This book uses Africa as an illuminating case study in the principles of ecology, and I used the opportunity of the Foreword to think about the relationship between ecology and natural selection. This could be seen as a continuation of my argument in the afterword to the previous section.
In this book, and elsewhere, I have been unkind to a view favoured by some social anthropologists, the 'cultural relativism' that acknowledges the equal status of many kinds of truth, scientific truth having no privileged rank among them. If ever I could be converted to some form of relativism, it might be after reading Elspeth Huxley's remarkable epic of Kenya, Red Strangers. Out of the Soul of Africa (6. 2) is the Foreword to the new paperback edition of her novel. I wrote an article for the Financial Times, pointing out that Red Strangers had been out of print for years and challenging any publisher to do something about it. The admirable Penguin did, and they reprinted my article as the Foreword.
I am now waiting for a scholar of literature to explain to me why Red Strangers is not rated one of the great novels of the twentieth century, the equal of a John Steinbeck except that Elspeth Huxley's imagery is Kikuyu rather than American.
Run like the eland . . . Run, warriors, with feet like arrows and the hearts of lions; the lives and wealth of your fathers are yours to save . . . Their thighs were straight as saplings, their features sharp as axes, their skins lighter than honey. His limbs began to quiver like the wings of a sunbird when its beak sucks honey . . .
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It is a virtuoso feat of identification with another culture. Not only does she succeed in getting herself inside a Kikuyu skin, she achieves the same feat for the reader. And she makes you cry.
I am slightly ashamed to admit that another book that brings me close to tears - of joy this time - is a children's book. Or is it a very grown-up book which happens to be written by children? It is hard to decide, which is part of its charm and also probably why it has been unaccountably ignored by book reviews editors - they just didn't know which shelf to put it on. The Lion Children is about a family of children who are English, but whose home is a set of tents in Botswana, where they radio-track wild lions and are schooled entirely by their mother in the bush. They have written a book about their utterly extraordinary life. Never mind whether there is a conventionally labelled shelf to put it on, just read it. I Speak of Africa and Golden Joys (6. 3), my Foreword, is reproduced here.
Last in this section is a travel piece, which again takes up the two themes of Africa as our ancestral home and Africa as my personal birthplace and weaves them together in an autobiographical story of travel and personal inspiration. The title was changed by the Sunday Times to 'All Our Yesterdays', but Macbeth's world-weariness is exactly opposite to the mood of my piece, so I am reverting to my original title, Heroes and Ancestors (6. 4). Now that I think about it, Heroes and Ancestors would have made another fine title for this whole collection.
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6 Ecology of Genes137
Foreword to Pyramids of Life
by Harvey Croze and John Reader
Africa was my personal cradle. But I left when I was seven, too young to appreciate - indeed the fact was not then known - that Africa is also humanity's cradle. The fossils of our species' formative years are all from Africa, and molecular evidence suggests that the ancestors of all today's peoples stayed there until as recently as the last hundred thousand years or so. We have Africa in our blood and Africa has our bones. We are all Africans.
This alone makes the African ecosystem an object of singular fascination. It is the community that shaped us, the commonwealth of animals and plants in which we served our ecological apprenticeship. But even if it were not our home continent Africa would captivate us, as perhaps the last great refugium of Pleistocene ecologies. If you want a late glimpse of the Garden of Eden, forget Tigris and Euphrates and the dawn of agriculture. Go instead to the Serengeti or the Kalahari. Forget the Arcadia of the Greeks and the dreamtime of the outback, they are so recent. Whatever may have come down the mountain at Olympus or Sinai, or even Ayers Rock, look instead to Kilimanjaro, or down the Rift Valley towards the High Veldt. There is where we were designed to flourish.
The 'design' of all living things and their organs is, of course, an illusion; an exceedingly powerful illusion, fabricated by a suitably power- ful process, Darwinian natural selection. There is a second illusion of design in nature, less compelling but still appealing, and it is in danger of being mistaken for the first. This is the apparent design of ecosystems. Where bodies have parts that intricately harmonize and regulate to keep them alive, ecosystems have species that appear to do something similar at a higher level. There are the primary producers that convert raw solar energy into a form that others can use. There are the herbivores that consume them to use it, and then make a tithe of it available for carnivores and so on up the food chain - pyramid, rather,
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for the laws of thermodynamics rule that only a tenth of each level's energy shall make it to the level above. Finally, there are scavengers that recycle the waste products to make them available again, and in the process clean up the world and stop it becoming a tip. Everything fits with everything else like jigsaw pieces meshing in a huge multi- dimensional puzzle, and - as the cliche goes - we meddle with the parts at the risk of destroying a priceless whole.
The temptation is to think that this second illusion is crafted by the same kind of process as the first: by a version of Darwinian selection, but at a higher level. According to this erroneous view, the ecosystems that survive are the ones whose parts - species - harmonize, just as the organisms that survive in conventional Darwinism are the ones whose parts - organs and cells - work harmoniously for their survival. I believe that this theory is false. Ecosystems, like organisms, do indeed seem harmoniously designed; and the appearance of design is indeed an illusion. But there the resemblance ends. It is a different kind of illusion, brought about by a different process. The best ecologists, such as Croze and Reader, understand this.
Darwinism enters into the process, but it does not jump levels. Genes still survive, or fail to survive, within the gene pools of species, by virtue of their effects upon the survival and reproduction of the individual organisms that contain them. The illusion of harmony at a higher level is an indirect consequence of differential individual reproduction. Within any one species of animals or plants, the individuals that survive best are the ones that can exploit the other animals and plants, bacteria and fungi that are already flourishing in the environment. As Adam Smith understood long ago, an illusion of harmony and real efficiency will emerge in an economy dominated by self-interest at a lower level. A well balanced ecosystem is an economy, not an adaptation.
Plants flourish for their own good, not for the good of herbivores. But because plants flourish, a niche for herbivores opens up, and they fill it. Grasses are said to benefit from being grazed. The truth is more interest- ing. No individual plant benefits from being grazed per se. But a plant that suffers only slightly when it is grazed outcompetes a rival plant that suffers more. So successful grasses have benefited indirectly from the presence of grazers. And of course grazers benefit from the presence of grasses. Grasslands therefore build up as harmonious communities of relatively compatible grasses and grazers. They seem to cooperate. In a sense they do, but it is a modest sense that must be cautiously under- stood and judiciously understated. The same is true of the other African communities expounded by Croze and Reader.
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? I have said that the illusion of harmony at the ecosystem level is its own kind of illusion, different from, and emphatically not to be confused with, the Darwinian illusion that produces each efficiently working body. But a closer look reveals that there is a similarity after all, one that goes deeper than the - admittedly interesting and more commonly stated - observation that an animal can also be seen as a community of symbiotic bacteria. Mainstream Darwinian selection is the differential survival of genes within gene pools. Genes survive if they build bodies that flourish in their normal environment. But the normal environ- ment of a gene importantly includes the other genes (strictly, their consequences) in the gene pool of the species. Natural selection therefore favours those genes that cooperate harmoniously in the joint enterprise of building bodies within the species. I have called the genes 'selfish cooperators'. There turns out to be, after all, an affinity between the harmony of a body and the harmony of an ecosystem. There is an ecology of genes.
ECOLOGY OF GENES
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Out of the Soul of Africa138 Foreword to Red Strangers by Elspeth Huxley
Elspeth Huxley died in 1997 at the age of 90. Best known for her vivid African memoirs, she was also a considerable novelist who, in Red Strangers, achieved a scale that could fairly be called epic. It is the saga of a Kikuyu family spanning four generations, beginning before the coming to Kenya of the British ('red' strangers because sunburned), and ending with the birth of a new baby girl, christened Aeroplane by her father ('His wife, he thought, would never be able to pronounce such a difficult word; but educated people would know, and understand'). Its 400 pages are gripping, moving, historically and anthropologically illuminating, humanistically mind-opening . . . and, lamentably, out of print. *
I had an unrealized youthful ambition to write a science fiction novel. It would follow an expedition to, say, Mars, but seen through the eyes (or whatever passed for eyes) of the native inhabitants. I wanted to manoeuvre my readers into an acceptance of Martian ways so compre- hensive that they would see the invading humans as strange and foreign aliens. It is Elspeth Huxley's extraordinary achievement in the first half of Red Strangers to immerse her readers so thoroughly in Kikuyu ways and thought that, when the British finally appear on the scene, everything about them seems to us alien, occasionally downright ridiculous, though usually to be viewed with indulgent tolerance. It is the same indulgent amusement, indeed, as I remember we bestowed upon Africans during my own colonial childhood.
Mrs Huxley, in effect, skilfully transforms her readers into Kikuyu, opening our eyes to see Europeans, and their customs, as we have never seen them before. We become used to an economy pegged to the Goat Standard, so when coins (first rupees and then shillings) are introduced,
*No longer!
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? we marvel at the absurdity of a currency that does not automatically accrue with each breeding season. We come to accept a world in which every event has a supernatural, magical interpretation, and feel personally swindled when the statement, 'The rupees that I pay you can later be changed into goats', turns out to be literally untrue. When Kichui (all white men are referred to by their Kikuyu nicknames) gives orders that hisfieldsshould be manured, we realize that he is mad. Why else would a man try to lay a curse upon his own cattle? 'Matu could not believe his ears. To bury the dung of a cow was to bring death upon it, just as death, or at any rate severe sickness, would come to a man whose excreta were covered with earth . . . He refused emphatically to obey the order. ' And, such is Elspeth Huxley's skill, even I, despising as I do the fashionable nostrums of 'cultural relativism', find myself endorsing Matu's sturdy good sense.
We are led to marvel at the absurdity of European justice, which seems to care which of two brothers committed a murder:
. . . what does it matter? Are not Muthengi and I brothers? Whichever it was that held the sword, our father Waseru and other members of our clan must still pay the blood-price.
Unaccountably, there is no blood price, and Matu, having cheerfully confessed to Muthengi's crime, goes to prison, where he leads 'a strange, comfortless life whose purpose he could not divine'. Eventually he is released. He has served his time but, since he didn't realize he was doing time, the event is of no significance. On returning to his own village, far from being disgraced, he has gained prestige from his sojourn with the mysterious strangers, who obviously regard him highly enough to invite him to live in their own place.
The novel takes us through episodes that we recognize as if from a great distance; through the First World War and the ravages of the subsequent Spanish flu, through smallpox epidemics and worldwide economic recession; and we never once are told in European terms that this is what is going on. We see all through Kikuyu eyes. The Germans are just another white tribe, and when the war ends we find ourselves wondering where are the plundered cattle that the victors ought to be driving home. What else, after all, is warfare for?
Ever since borrowing Red Strangers from the library, I have been on a ceaseless quest to acquire a copy of my own. It has been my routine first question on every visit to second-hand book fairs. Finally, I tracked down two old American copies simultaneously on the Internet. After so
OUT OF THE SOUL OP AFRICA
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many years of restless searching, I could not resist buying both. So now, if any reputable publisher sincerely wants to look at Red Strangers with a view to bringing out a new edition,* I will gladly make available one of my hard-won copies. Nothing will part me from the other one.
*This was first published in the Financial Times. I am delighted to say that Penguin Books rose to this challenge and published the book, using my Financial Times article, here reproduced again, as the Foreword.
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I Speak of Africa and Golden Joys1
Foreword to The Lion Children
by Angus, Maisie and Travers McNeice
This is an astonishing book, by an even more astonishing trio of children. It's hard to describe: you have to read it, and once you start reading you can't stop. Think of Swallows and Amazons, except that this story is true and it all happens far from the comfort of England. Think
of The Lion, the Witch and the Wardrobe, except that the Lion Children need no magic wardrobe to pass through; no fake world of wonder. The real Africa, humanity's cradle, is more magical than anything C. S. Lewis could dream up. And, while they have no witch, these young authors do have a most remarkable mother. More of her in a moment.
Travers, Angus, Maisie and family have lived under canvas for almost as long as their little brother Oakley (think of fust William) can remember. All three of them have been driving Land Rovers ever since their feet could reach the pedals, and changing tyres (frequently) for as long as they've been strong enough to lift them. * They are self-sufficient and trustworthy far beyond their years, yet not in that disagreeable sense of being streetwise and fly. Field Marshall Montgomery once described Mao Tse Tung as the sort of man you could go into the jungle with. Well, I'm not sure I'd go with Mao Tse Tung into Hyde Park, but I would unhesitatingly go into the jungle with Travers, Angus and Maisie, and no adult companions at all. No gun, just quick-witted young people with clear eyes, fast reflexes and most of a lifetime (albeit a short one) of African know-how. I don't know what to do if I meet an elephant. They do. I'm terrified of puff adders, mambas and scorpions. They take them in their stride. At the same time, dependable and strong as they are, they still bubble with the innocence and charm of youth. This is still Swallows and Amazons, still an idyll, the sort of childhood that for most of us exists only in dreams and idealized misrememberings, 'the land of lost content'. Yet it is firmly in the real world. These innocents have seen favourite lions brutally killed, have
*Travers, Angus and Maisie were aged 16, 14 and 12 when they finished the book.
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rapped out reports of such tragedies in the dispassionate argot of the radio link, have assisted at the subsequent postmortems.
This accomplished book is entirely the work of its young authors, but it isn't hard to guess the source of their ability to do it - their imagina- tion, their enterprise, their unorthodoxy, their adventurous spirit. My wife and I first met Kate Nicholls, their mother, in 1992 when she was living in the Cotswolds, pregnant with Oakley, commuting to study in Oxford libraries. A successful actress, she had become disillusioned with the stage and developed, in her late thirties, a passion (passion is the story of her life) for the science of evolution. Kate doesn't do anything by halves and, for her, an interest in evolution meant deep immersion in libraries, digging up the original research literature. With only minimal guidance from me in what became a series of informal tutorials, her reading transformed her into something of a scholarly authority on Darwinian theory. Her eventual decision to pull up roots and head for Botswana, where Darwinism can be daily witnessed in practice, seemed entirely in character: a natural, if unconventional, extension of the same scholarly quest. Her children, one can't help feeling, have a pretty fortunate inheritance, as well as an almost unique environment in which to realize it.
They also have to thank their mother for their education, and this is perhaps the most surprising aspect of their life. Quite soon after arriving in Botswana, Kate decided to teach them herself. A brave decision, I think I would have counselled against it. But I would have been wrong. Although all their schooling is done in camp, they keep proper terms, have challenging homework assignments and work towards inter- nationally accredited exams. Kate gets good results by standard educational certifications, while at the same time tending, indeed enhancing, the natural sense of wonder that normal children too often lose during their teens. I don't think any reader of these pages could fail to judge her unorthodox School in the Bush a brilliant success. * The proof lies in the book, for, to repeat, the children, and they alone, wrote it. All three authors show themselves to be excellent writers: sensitive, literate, articulate, intelligent and creative.
Kate's choice of Botswana rather than anywhere else in Africa was for- tuitous. It eventually led to her meeting Pieter Kat. And of course the lions - wild lions, living and dying in the world for which the natural selection of their ancestors had prepared them. Pieter is the ideal step-
? Further testimony to the success of Kate's bush school is that Travers and Angus have both been accepted by the (first-class) universities of their choice, UC Santa Barbara and Stanford, respectively.
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? father for her children, and these young scientists have in turn become an indispensable part of the lion research and conservation project.
It wasn't till last year that my family and I finally visited the camp. The experience was unforgettable, and I can testify to the picture painted in The Lion Children. It really is just like that: more wonderful than mad, but a bit of both. My daughter Juliet went out ahead, part of a large invasion of young visitors who soon picked up the enthusiasm of the resident family. On Juliet's first full day in Africa, Travers took her out in a Land Rover, tracking radio-collared lions. When we received Juliet's letter home, brimming with excitement at such an initiation, I relayed the story to her grandmother, who interrupted me with panic in her voice: 'Plus, of course, at least two armed African rangers?
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to service the arms race. And in any case the improvement in equip-
ment will normally be progressive. Another kind of positive feedback in evolution, if R. A. Fisher and his followers are right, results from sexual selection. Once again, progressive evolution is the expected consequence.
Progressive increase in morphological complexity is to be expected only in taxa whose way of life benefits from morphological complexity. Progressive increase in brain size is to be expected only in animals where braininess is an advantage. This may, for all I know, constitute a minority of lineages. But what I do insist is that in a majority of evolutionary lineages there will be progressive evolution towards some- thing. It won't, however, be the same thing in different lineages (this was the point about swifts and elephants). And there is no general reason to expect a majority of lineages to progress in the directions pioneered by our human line.
But have I now defined progress so generally as to make it a blandly useless word? I don't think so. To say that the evolution of the verte- brate eye was progressive is to say something quite strong and quite important. If you could lay out all the intermediate ancestors in chronological order you'd find that, first, for a majority of dimensions
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of measurement, the changes would be transitive over the whole sequence. That is, if A is ancestral to B which is ancestral to C, the direction of change from A to B is likely to be the same as the direction of change from B to C. Second, the number of successive steps over which progress is seen is likely to be large: the transitive series extends beyond A, B and C, far down the alphabet. Third, an engineer would judge the performance to have improved over the sequence. Fourth, the number of separate features combining and conspiring to improve performance would increase. Finally, this kind of progress really matters because it is the key to answering the Hoyle challenge. There will be exceptional reversals, for instance in the evolution of blind cave fish, where eyes degenerate because they are not used and are costly to make. And there will doubtless be periods of stasis where there is no evolution at all, progressive or otherwise.
To conclude this point, Gould is wrong to say that the appearance of progress in evolution is a statistical illusion. It does not result just from a change in variance as a baseball-style artefact. To be sure, complexity, braininess and other particular qualities dear to the human ego should not necessarily be expected to increase progressively in a majority of lineages - though it would be interesting if they did: the investigations
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of McShea, Jerison and others are not a waste of time. But if you
define progress less chauvinistically - if you let the animals bring their own definition - you will find progress, in a genuinely interesting sense of the word, nearly everywhere.
Now it is important to stress that, on this adaptationist view (unlike the 'evolution of evolvability' view to be discussed shortly), progressive evolution is to be expected only on the short to medium term. Coevolutionary arms races may last for millions of years, but probably not hundreds of millions. Over the very long timescale, asteroids and other catastrophes bring evolution to a dead stop, major taxa and entire radiations go extinct. Ecological vacuums are created, to befilledby new adaptive radiations driven by new ranges of arms races. The several arms races between carnivorous dinosaurs and their prey were later mirrored by a succession of analogous arms races between carnivorous mammals and their prey. Each of these successive and separate arms races powered sequences of evolution which were progressive in my sense. But there was no global progress over the hundreds of millions of years, only a sawtooth succession of small progresses terminated by extinctions. Nonetheless, the ramp phase of each sawtooth was properly and significantly progressive.
Ironically for such an eloquent foe of progress, Gould flirts with the idea that evolution itself changes over the long haul, but he puts it in a topsy-turvy way which has undoubtedly been widely misleading. It is
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more fully expounded in Wonderful Life but reprised in the present book. For Gould, evolution in the Cambrian was a different kind of process from evolution today. The Cambrian was a period of evolution- ary 'experiment', evolutionary 'trial and error', evolutionary 'false starts'. It was a period of 'explosive' invention, before evolution stabilized into the humdrum process we see today. It was the fertile time when all the great 'fundamental body plans' were invented. Nowadays, evolution just tinkers with old body plans. Back in the Cambrian, new phyla and new classes arose. Nowadays we only get new species!
This may be a slight caricature of Gould's own considered position,
but there is no doubt that the many American nonspecialists who
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unfortunately, as Maynard Smith wickedly observes, get their
evolutionary knowledge almost entirely from Gould, have been deeply misled. Admittedly, what follows is an extreme example, but Daniel Dennett has recounted a conversation with a philosopher colleague who read Wonderful Life as arguing that the Cambrian phyla did not have a common ancestor - that they had sprung up as independently initiated life forms! When Dennett assured him that this was not Gould's claim, his colleague's response was, 'Well then, what is all the fuss about? '
Even some professional evolutionists have been inspired by Gould's rhetoric into committing some pretty remarkable solecisms. Leakey and
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Lewin's The Sixth Extinction is an excellent book except for its Chapter
3, 'The Mainspring of Evolution', which is avowedly heavily influenced by Gould. The following quotations from that chapter could hardly be more embarrassingly explicit:
Why haven't new animal body plans continued to crawl out of the evolutionary cauldron during the past hundreds of millions of years?
In early Cambrian times, innovations at the phylum level survived because they faced little competition.
Below the level of the family, the Cambrian explosion produced relatively few species, whereas in the post-Permian a tremendous species diversity burgeoned. Above family level however, the post-Permian radiation faltered, with few new classes and no new phyla being generated. Evidently, the mainspring of evolution operated in both periods, but it propelled greater extreme experimentation in the Cambrian than in the post-Permian, and greater variations on existing themes in the post-Permian.
Hence, evolution in Cambrian organisms could take bigger leaps, including phylum-level leaps, while later on it would be more constrained, making only modest jumps, up to the class level.
It is as though a gardener looked at an old oak tree and remarked,
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wonderingly: 'Isn't it strange that no major new boughs have appeared on this tree recently. These days, all the new growth appears to be at the twig level! '
As it happens, molecular clock evidence indicates that the 'Cambrian Explosion' may never have happened. Far from the major phyla diverging from a point at the beginning of the Cambrian, Wray,
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Levinton and Shapiro present evidence that the common ancestors of
the major phyla are staggered through hundreds of millions of years back in the Precambrian. But never mind that. That is not the point I want to make. Even if there really was a Cambrian explosion such that all the major phyla diverged during a ten million-year period, this is no reason to think that Cambrian evolution was a qualitatively special kind of super-jumpy process. Bauplane don't drop out of a clear Platonic sky, they evolve step by step from predecessors, and they do so (I bet, and so would Gould if explicitly challenged) under approximately the same Darwinian rules as we see today.
'Phylum-level leaps' and 'modest jumps, up to the class level' are the sheerest nonsense. Jumps above the species level don't happen, and nobody who thinks about it for two minutes claims that they do. Even the great phyla, when they originally bifurcated one from another, were just pairs of new species, members of the same genus. Classes are species that diverged a very long time ago, and phyla are species that diverged an even longer time ago. Indeed it is a moot - and rather empty - question precisely when in the course of the step-by-step, gradual mutual divergence of, say, mollusc ancestors and annelid ancestors after the time when they were congeneric species, we should wish to say that the divergence had reached 'Bauplan' status. A good case could be made that The Bauplan is a myth, probably as pernicious as any of the myths that Stephen Gould has so ably combatted, but this one, in its modern form, is largely perpetuated by him.
I return, finally, to the 'evolution of evolvability' and a very real sense in which evolution itself may evolve, progressively, over a longer timescale than the individual ramps of the arms race sawtooth. Notwith- standing Gould's just scepticism over the tendency to label each era by its newest arrivals, there really is a good possibility that major innovations in embryological technique open up new vistas of evolutionary possibility and that these constitute genuinely progressive improvements. * The
*This is the idea that I dubbed 'The Evolution of Evolvability' (in C. Langton (ed. ), Artificial Life (Santa Fe, Addison Wesley, 1982)) and Maynard Smith and Szathmary wrote a book about 0. Maynard Smith and E. Szathmary, The Major Transitions in Evolution (Oxford, W. H. Freeman/Spektrum, 1995)).
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origin of the chromosome, of the bounded cell, of organized meiosis, diploidy and sex, of the eucaryotic cell, of multicellularity, of gastrulation, of molluscan torsion, of segmentation - each of these may have constituted a watershed event in the history of life. Not just in the normal Darwinian sense of assisting individuals to survive and reproduce, but watershed in the sense of boosting evolution itself in ways that seem entitled to the label progressive. It may well be that after, say, the invention of multicellularity, or the invention of segmentation, evolution was never the same again. In this sense there may be a one-way ratchet of progressive innovation in evolution.
For this reason over the long term, and because of the cumulative character of coevolutionary arms races over the shorter term, Gould's attempt to reduce all progress to a trivial, baseball-style artefact con- stitutes a surprising impoverishment, an uncharacteristic slight, an unwonted demeaning of the richness of evolutionary processes.
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mm* Mmi. W SI^b#
Unfinished Correspondence with a Darwinian Heavyweight
The following correspondence was never completed and now, sadly, it never can be.
9 December 2001
Stephen Jay Gould Harvard
Dear Steve
Recently I received an email from Phillip Johnson, founder of the so-called 'Intelligent Design' school of creationists, crowing in triumph because one of his colleagues, Jonathan Wells, had been invited to take part in a debate at Harvard. He included the text of his email on his 'Wedge of Truth' web site, in which he announced the Wells debate under the headline 'Wells Hits a Home Run at Harvard'.
http://www. arn. org/docs/pjweekly/pj_weekly_011202. htm
The 'Home Run' turns out to be NOT a resounding success by Wells in convincing the audience, NOR any kind of besting of his opponent (Stephen Palumbi, who tells me he agreed to take part, with great reluctance, only because somebody at Harvard had ALREADY invited Wells and it was too late to do anything about that). There is no suggestion that Wells did well in the debate, nor even any obvious interest in whether he did. No, the 'Home Run' was simply and solely the matter of being invited by Harvard in the first place. These people have no hope of convincing reputable scientists by their ridiculous arguments. Instead, what they seek is the oxygen of respectability. We give them this oxygen by the mere act of ENGAGING with them at all. They don't mind being beaten in argument. What matters is that we give them recognition by bothering to argue with them in public.
You convinced me of this years ago when I phoned you up (you have probably
forgotten this) to ask your advice when I was invited to debate Duane P Gish. 218
? Ever since that phone call, I have repeatedly cited you and refused to debate these people, not because I am afraid of 'losing' the debate, but because, as you said, just to appear on a platform with them is to lend them the respectability they crave. Whatever might be the outcome of the debate, the mere fact that it is staged at all suggests to ignorant bystanders that there must be something worth debating, on something like equal terms.
First, I am interested to know whether you still hold to this view, as I do. Second, I am proposing that you might consider uniting with me (no need to involve others) in signing a short letter, say to the New York Review of Books, explaining publicly why we do not debate creationists (including the 'Intelligent Design' euphemism for creationists) and encouraging other evolutionary biologists to follow suit.
Such a letter would have great impact precisely because there have been widely publicised differences, and even animosities, between us (differences which creationists, with extreme intellectual dishonesty, have not hesitated to exploit). And I would not suggest writing a long disquisition on the technical differences which remain between us. That would only confuse the issue, make it harder to agree on a final draft, and lessen the impact. I wouldn't even mention our differences. I suggest a brief letter to the editor, explaining why we do not engage with 'intelligent design' or any other species of creationists, and offering our letter as a model for others to cite in refusing such invitations in the future. We both have better things to do with our time than give it over to such nonsense. Having just reached my sixtieth birthday (we are almost exactly the same age) I feel this keenly.
Steve replied on 11th December 2001, a warm and friendly email enthusiastically agreeing that a joint letter was an excellent idea, and saying that he'd be delighted to join me as the sole other signatory. He agreed that the New York Review of Books might well be the best place, and proposed that I should write a first draft. I reproduce that here, exactly as I sent it to him for his approval.
14 December 2001
Dear Editor
Like any flourishing science, the study of evolution has its internal controversies, as we both know. But no qualified scientist doubts that evolution is a fact, in the ordinarily accepted sense in which it is a fact that the Earth orbits the Sun. It is a fact that human beings are cousins
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to monkeys, kangaroos, jellyfish and bacteria. No reputable biologist doubts this. Nor do reputable theologians, from the Pope on. Unfortunately, many lay Americans do, including some frighteningly influential, powerful and, above all, well-financed ones.
We are continually invited to engage in public debates against creationists, including latter-day creationists disguised under the euphemism 'Intelligent Design Theorists'. We always refuse, for one overriding reason. If we may be allowed to spell this reason out publicly, we hope our letter may be helpful to other evolutionary scientists plagued by similar invitations.
The question of who would 'win' such a debate is not at issue. Winning is not what these people realistically aspire to. The coup they seek is simply the recognition of being allowed to share a platform with a real scientist in thefirstplace. This will suggest to innocent bystanders that there must be material here that is genuinely worth debating, on something like equal terms.
At the moment of writing, the leading 'Intelligent Design' website
reports a debate at Harvard under the banner headline 'Wells Hits a
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Home Run at Harvard'. Jonathan Wells is a creationist, incidentally a
long-time devotee of the Unification Church (the 'Moonies'). * He had a debate last month against Stephen Palumbi, Professor of Biology at Harvard University. 'Home Run' might seem to suggest that Reverends (sic) Wells scored some kind of victory over Professor Palumbi. Or at least that he made powerful points and his speech was well received. No such claim is made. It doesn't even seem to be of interest.
The 'Home Run' turns out to be simply the public demonstration at Harvard that, in the words of the website's author, Phillip Johnson, 'This is the sort of debate that is now occurring in universities. ' There was a victory, but it occurred long before the debate itself. The creationist scored his home run at the moment the invitation from Harvard landed on his doormat. It came, by the way, not from any biological, or indeed scientific, department, but from the Institute of Politics.
Phillip Johnson himself, founding father of the 'Intelligent Design'
''Darwinism: Why I went for a Second Ph. D. ' is Jonathan Wells' own testimony on the turning point of his life: 'Father's words, my studies, and my prayers convinced me that I should devote my life to destroying Darwinism, just as many of my fellow Unificationists had already devoted their lives to destroying Marxism. When Father chose me (along with about a dozen other seminary graduates) to enter a Ph. D. program in 1978, I welcomed the opportunity to prepare myself for battle. ' ('Father', of course, is the Moonies' name for Rev Moon himself). http://www. tparents. org/LibraryAJnincation/Talks/Wells/DARWIN. htm.
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? movement (not a biologist, nor a scientist of any kind, but a lawyer who became a mid-life born-again Christian), wrote, in a letter of 6 April 2001, which he copied to one of us:
It isn't worth my while to debate every ambitious Darwinist who wants to try his hand at ridiculing the opposition, so my general policy is that Darwinists have to put a significant figure at risk before I will agree to a debate. That means specifically Dawkins or Gould, or someone of like stature and public visibility.
Well, we can condescend too, and we have the advantage that evolutionary scientists don't need the publicity such debates can bring. In the unlikely event that a significant argument should ever emerge from the ranks of creationism/'intelligent design', we will be happy to debate it. Meanwhile, we shall cultivate our evolutionary gardens, occasionally engaging in the more exacting and worthwhile task of debating each other. What we shall not do is abet creationists in their disreputable quest for free publicity and unearned academic respectability.
In all humility, we offer these thoughts to our colleagues who receive similar invitations to debate.
Unfortunately, Steve never got around to revising the letter, which therefore lacks the stylish panache which his dexterous touch would have lent it. I received one further email, apologizing for the delay and hoping to deal with the matter soon. The subsequent silence, I now realize, coincided with his last illness. I therefore offer my draft, imperfect as it is, in the hope that it may go some way towards conveying the message which I originally learned from him many years ago. It is my sincere hope that he would have approved the content of the letter, but of course I cannot be sure.
To close this section on a note of such concord may seem puzzling.
Given that Steve was as much a neo-Darwinist as I am, what did we dis-
agree about? The major disagreement emerges clearly out of his last big
5
book, The Structure of Evolutionary Theory,TM which I had no opportunity
to see until after his death. It is appropriate, therefore, to spell out that issue here, and it also, as it happens, forms a natural bridge to the next essay. The question under dispute is this: what is the role of genes in evolution? Is it, to use Gould's phrase, 'book-keeping or causation'?
Gould saw natural selection as operating on many levels in the hierarchy of life. Indeed it may, after a fashion, but I believe that such selection can have evolutionary consequences only when the entities selected consist of 'replicators'. A replicator is a unit of coded information, of high fidelity but occasionally mutable, with some causal power over its own fate. Genes are
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such entities. So, in principle, are memes, but they are not under discus- sion here. Biological natural selection, at whatever level we may see it, results in evolutionary effects only insofar as it gives rise to changes in gene frequencies in gene pools. Gould, however, saw genes only as 'book- keepers', passively tracking the changes going on at other levels. In my view, whatever else genes are, they must be more than book-keepers, otherwise natural selection cannot work. If a genetic change has no causal influence on bodies, or at least on something that natural selection can 'see', natural selection cannot favour or disfavour it. No evolutionary change will result.
Gould and I would agree that genes can be seen as a book in which is
written the evolutionary history of a species. In Unweaving the Rainbow I
called it 'The Genetic Book of the Dead'. But the book is written via the
natural selection of randomly varying genes, chosen by virtue of their
causal influence on bodies. Book-keeping is precisely the wrong metaphor,
because it reverses the causal arrow, almost in Lamarckian fashion, and
makes the genes passive recorders. I dealt with this in 1982 (The Extended
Phenotype) in my distinction between 'active replicators' and 'passive
replicators'. The point is also explained in David Barash's superb review of
136 Gould's book.
Book-keeping is perversely - and characteristically - a valuable meta- phor precisely because it is so diametrically back to front. Not for the first time, the characteristic vividness and clarity of a Gouldian metaphor helps us to see vividly and clearly what is wrong with the Gouldian message - and how it needs to be reversed in order to get at the truth.
I hope this brief note will not be seen as taking advantage to get the last word. The Structure of Evolutionary Theory is such a massively powerful last word, it will keep us all busy replying to it for years. What a brilliant way
for a scholar to go. I shall miss him.
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THERE IS ALL AFRICA AND HER PRODIGIES IN US
I am one of those (it includes most people who have ever spent time south of the Sahara) who think of Africa as a place of enchantment. For me it stems from faint but haunting childhood memories, coupled with the mature understanding that Africa is our ancestral home.
These themes recur throughout this section, and they introduce Ecology of Genes (6. 1), my Foreword to Harvey Croze and John Reader's Pyramids of Life. This book uses Africa as an illuminating case study in the principles of ecology, and I used the opportunity of the Foreword to think about the relationship between ecology and natural selection. This could be seen as a continuation of my argument in the afterword to the previous section.
In this book, and elsewhere, I have been unkind to a view favoured by some social anthropologists, the 'cultural relativism' that acknowledges the equal status of many kinds of truth, scientific truth having no privileged rank among them. If ever I could be converted to some form of relativism, it might be after reading Elspeth Huxley's remarkable epic of Kenya, Red Strangers. Out of the Soul of Africa (6. 2) is the Foreword to the new paperback edition of her novel. I wrote an article for the Financial Times, pointing out that Red Strangers had been out of print for years and challenging any publisher to do something about it. The admirable Penguin did, and they reprinted my article as the Foreword.
I am now waiting for a scholar of literature to explain to me why Red Strangers is not rated one of the great novels of the twentieth century, the equal of a John Steinbeck except that Elspeth Huxley's imagery is Kikuyu rather than American.
Run like the eland . . . Run, warriors, with feet like arrows and the hearts of lions; the lives and wealth of your fathers are yours to save . . . Their thighs were straight as saplings, their features sharp as axes, their skins lighter than honey. His limbs began to quiver like the wings of a sunbird when its beak sucks honey . . .
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It is a virtuoso feat of identification with another culture. Not only does she succeed in getting herself inside a Kikuyu skin, she achieves the same feat for the reader. And she makes you cry.
I am slightly ashamed to admit that another book that brings me close to tears - of joy this time - is a children's book. Or is it a very grown-up book which happens to be written by children? It is hard to decide, which is part of its charm and also probably why it has been unaccountably ignored by book reviews editors - they just didn't know which shelf to put it on. The Lion Children is about a family of children who are English, but whose home is a set of tents in Botswana, where they radio-track wild lions and are schooled entirely by their mother in the bush. They have written a book about their utterly extraordinary life. Never mind whether there is a conventionally labelled shelf to put it on, just read it. I Speak of Africa and Golden Joys (6. 3), my Foreword, is reproduced here.
Last in this section is a travel piece, which again takes up the two themes of Africa as our ancestral home and Africa as my personal birthplace and weaves them together in an autobiographical story of travel and personal inspiration. The title was changed by the Sunday Times to 'All Our Yesterdays', but Macbeth's world-weariness is exactly opposite to the mood of my piece, so I am reverting to my original title, Heroes and Ancestors (6. 4). Now that I think about it, Heroes and Ancestors would have made another fine title for this whole collection.
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6 Ecology of Genes137
Foreword to Pyramids of Life
by Harvey Croze and John Reader
Africa was my personal cradle. But I left when I was seven, too young to appreciate - indeed the fact was not then known - that Africa is also humanity's cradle. The fossils of our species' formative years are all from Africa, and molecular evidence suggests that the ancestors of all today's peoples stayed there until as recently as the last hundred thousand years or so. We have Africa in our blood and Africa has our bones. We are all Africans.
This alone makes the African ecosystem an object of singular fascination. It is the community that shaped us, the commonwealth of animals and plants in which we served our ecological apprenticeship. But even if it were not our home continent Africa would captivate us, as perhaps the last great refugium of Pleistocene ecologies. If you want a late glimpse of the Garden of Eden, forget Tigris and Euphrates and the dawn of agriculture. Go instead to the Serengeti or the Kalahari. Forget the Arcadia of the Greeks and the dreamtime of the outback, they are so recent. Whatever may have come down the mountain at Olympus or Sinai, or even Ayers Rock, look instead to Kilimanjaro, or down the Rift Valley towards the High Veldt. There is where we were designed to flourish.
The 'design' of all living things and their organs is, of course, an illusion; an exceedingly powerful illusion, fabricated by a suitably power- ful process, Darwinian natural selection. There is a second illusion of design in nature, less compelling but still appealing, and it is in danger of being mistaken for the first. This is the apparent design of ecosystems. Where bodies have parts that intricately harmonize and regulate to keep them alive, ecosystems have species that appear to do something similar at a higher level. There are the primary producers that convert raw solar energy into a form that others can use. There are the herbivores that consume them to use it, and then make a tithe of it available for carnivores and so on up the food chain - pyramid, rather,
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for the laws of thermodynamics rule that only a tenth of each level's energy shall make it to the level above. Finally, there are scavengers that recycle the waste products to make them available again, and in the process clean up the world and stop it becoming a tip. Everything fits with everything else like jigsaw pieces meshing in a huge multi- dimensional puzzle, and - as the cliche goes - we meddle with the parts at the risk of destroying a priceless whole.
The temptation is to think that this second illusion is crafted by the same kind of process as the first: by a version of Darwinian selection, but at a higher level. According to this erroneous view, the ecosystems that survive are the ones whose parts - species - harmonize, just as the organisms that survive in conventional Darwinism are the ones whose parts - organs and cells - work harmoniously for their survival. I believe that this theory is false. Ecosystems, like organisms, do indeed seem harmoniously designed; and the appearance of design is indeed an illusion. But there the resemblance ends. It is a different kind of illusion, brought about by a different process. The best ecologists, such as Croze and Reader, understand this.
Darwinism enters into the process, but it does not jump levels. Genes still survive, or fail to survive, within the gene pools of species, by virtue of their effects upon the survival and reproduction of the individual organisms that contain them. The illusion of harmony at a higher level is an indirect consequence of differential individual reproduction. Within any one species of animals or plants, the individuals that survive best are the ones that can exploit the other animals and plants, bacteria and fungi that are already flourishing in the environment. As Adam Smith understood long ago, an illusion of harmony and real efficiency will emerge in an economy dominated by self-interest at a lower level. A well balanced ecosystem is an economy, not an adaptation.
Plants flourish for their own good, not for the good of herbivores. But because plants flourish, a niche for herbivores opens up, and they fill it. Grasses are said to benefit from being grazed. The truth is more interest- ing. No individual plant benefits from being grazed per se. But a plant that suffers only slightly when it is grazed outcompetes a rival plant that suffers more. So successful grasses have benefited indirectly from the presence of grazers. And of course grazers benefit from the presence of grasses. Grasslands therefore build up as harmonious communities of relatively compatible grasses and grazers. They seem to cooperate. In a sense they do, but it is a modest sense that must be cautiously under- stood and judiciously understated. The same is true of the other African communities expounded by Croze and Reader.
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? I have said that the illusion of harmony at the ecosystem level is its own kind of illusion, different from, and emphatically not to be confused with, the Darwinian illusion that produces each efficiently working body. But a closer look reveals that there is a similarity after all, one that goes deeper than the - admittedly interesting and more commonly stated - observation that an animal can also be seen as a community of symbiotic bacteria. Mainstream Darwinian selection is the differential survival of genes within gene pools. Genes survive if they build bodies that flourish in their normal environment. But the normal environ- ment of a gene importantly includes the other genes (strictly, their consequences) in the gene pool of the species. Natural selection therefore favours those genes that cooperate harmoniously in the joint enterprise of building bodies within the species. I have called the genes 'selfish cooperators'. There turns out to be, after all, an affinity between the harmony of a body and the harmony of an ecosystem. There is an ecology of genes.
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? 6. 2
Out of the Soul of Africa138 Foreword to Red Strangers by Elspeth Huxley
Elspeth Huxley died in 1997 at the age of 90. Best known for her vivid African memoirs, she was also a considerable novelist who, in Red Strangers, achieved a scale that could fairly be called epic. It is the saga of a Kikuyu family spanning four generations, beginning before the coming to Kenya of the British ('red' strangers because sunburned), and ending with the birth of a new baby girl, christened Aeroplane by her father ('His wife, he thought, would never be able to pronounce such a difficult word; but educated people would know, and understand'). Its 400 pages are gripping, moving, historically and anthropologically illuminating, humanistically mind-opening . . . and, lamentably, out of print. *
I had an unrealized youthful ambition to write a science fiction novel. It would follow an expedition to, say, Mars, but seen through the eyes (or whatever passed for eyes) of the native inhabitants. I wanted to manoeuvre my readers into an acceptance of Martian ways so compre- hensive that they would see the invading humans as strange and foreign aliens. It is Elspeth Huxley's extraordinary achievement in the first half of Red Strangers to immerse her readers so thoroughly in Kikuyu ways and thought that, when the British finally appear on the scene, everything about them seems to us alien, occasionally downright ridiculous, though usually to be viewed with indulgent tolerance. It is the same indulgent amusement, indeed, as I remember we bestowed upon Africans during my own colonial childhood.
Mrs Huxley, in effect, skilfully transforms her readers into Kikuyu, opening our eyes to see Europeans, and their customs, as we have never seen them before. We become used to an economy pegged to the Goat Standard, so when coins (first rupees and then shillings) are introduced,
*No longer!
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? we marvel at the absurdity of a currency that does not automatically accrue with each breeding season. We come to accept a world in which every event has a supernatural, magical interpretation, and feel personally swindled when the statement, 'The rupees that I pay you can later be changed into goats', turns out to be literally untrue. When Kichui (all white men are referred to by their Kikuyu nicknames) gives orders that hisfieldsshould be manured, we realize that he is mad. Why else would a man try to lay a curse upon his own cattle? 'Matu could not believe his ears. To bury the dung of a cow was to bring death upon it, just as death, or at any rate severe sickness, would come to a man whose excreta were covered with earth . . . He refused emphatically to obey the order. ' And, such is Elspeth Huxley's skill, even I, despising as I do the fashionable nostrums of 'cultural relativism', find myself endorsing Matu's sturdy good sense.
We are led to marvel at the absurdity of European justice, which seems to care which of two brothers committed a murder:
. . . what does it matter? Are not Muthengi and I brothers? Whichever it was that held the sword, our father Waseru and other members of our clan must still pay the blood-price.
Unaccountably, there is no blood price, and Matu, having cheerfully confessed to Muthengi's crime, goes to prison, where he leads 'a strange, comfortless life whose purpose he could not divine'. Eventually he is released. He has served his time but, since he didn't realize he was doing time, the event is of no significance. On returning to his own village, far from being disgraced, he has gained prestige from his sojourn with the mysterious strangers, who obviously regard him highly enough to invite him to live in their own place.
The novel takes us through episodes that we recognize as if from a great distance; through the First World War and the ravages of the subsequent Spanish flu, through smallpox epidemics and worldwide economic recession; and we never once are told in European terms that this is what is going on. We see all through Kikuyu eyes. The Germans are just another white tribe, and when the war ends we find ourselves wondering where are the plundered cattle that the victors ought to be driving home. What else, after all, is warfare for?
Ever since borrowing Red Strangers from the library, I have been on a ceaseless quest to acquire a copy of my own. It has been my routine first question on every visit to second-hand book fairs. Finally, I tracked down two old American copies simultaneously on the Internet. After so
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many years of restless searching, I could not resist buying both. So now, if any reputable publisher sincerely wants to look at Red Strangers with a view to bringing out a new edition,* I will gladly make available one of my hard-won copies. Nothing will part me from the other one.
*This was first published in the Financial Times. I am delighted to say that Penguin Books rose to this challenge and published the book, using my Financial Times article, here reproduced again, as the Foreword.
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I Speak of Africa and Golden Joys1
Foreword to The Lion Children
by Angus, Maisie and Travers McNeice
This is an astonishing book, by an even more astonishing trio of children. It's hard to describe: you have to read it, and once you start reading you can't stop. Think of Swallows and Amazons, except that this story is true and it all happens far from the comfort of England. Think
of The Lion, the Witch and the Wardrobe, except that the Lion Children need no magic wardrobe to pass through; no fake world of wonder. The real Africa, humanity's cradle, is more magical than anything C. S. Lewis could dream up. And, while they have no witch, these young authors do have a most remarkable mother. More of her in a moment.
Travers, Angus, Maisie and family have lived under canvas for almost as long as their little brother Oakley (think of fust William) can remember. All three of them have been driving Land Rovers ever since their feet could reach the pedals, and changing tyres (frequently) for as long as they've been strong enough to lift them. * They are self-sufficient and trustworthy far beyond their years, yet not in that disagreeable sense of being streetwise and fly. Field Marshall Montgomery once described Mao Tse Tung as the sort of man you could go into the jungle with. Well, I'm not sure I'd go with Mao Tse Tung into Hyde Park, but I would unhesitatingly go into the jungle with Travers, Angus and Maisie, and no adult companions at all. No gun, just quick-witted young people with clear eyes, fast reflexes and most of a lifetime (albeit a short one) of African know-how. I don't know what to do if I meet an elephant. They do. I'm terrified of puff adders, mambas and scorpions. They take them in their stride. At the same time, dependable and strong as they are, they still bubble with the innocence and charm of youth. This is still Swallows and Amazons, still an idyll, the sort of childhood that for most of us exists only in dreams and idealized misrememberings, 'the land of lost content'. Yet it is firmly in the real world. These innocents have seen favourite lions brutally killed, have
*Travers, Angus and Maisie were aged 16, 14 and 12 when they finished the book.
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rapped out reports of such tragedies in the dispassionate argot of the radio link, have assisted at the subsequent postmortems.
This accomplished book is entirely the work of its young authors, but it isn't hard to guess the source of their ability to do it - their imagina- tion, their enterprise, their unorthodoxy, their adventurous spirit. My wife and I first met Kate Nicholls, their mother, in 1992 when she was living in the Cotswolds, pregnant with Oakley, commuting to study in Oxford libraries. A successful actress, she had become disillusioned with the stage and developed, in her late thirties, a passion (passion is the story of her life) for the science of evolution. Kate doesn't do anything by halves and, for her, an interest in evolution meant deep immersion in libraries, digging up the original research literature. With only minimal guidance from me in what became a series of informal tutorials, her reading transformed her into something of a scholarly authority on Darwinian theory. Her eventual decision to pull up roots and head for Botswana, where Darwinism can be daily witnessed in practice, seemed entirely in character: a natural, if unconventional, extension of the same scholarly quest. Her children, one can't help feeling, have a pretty fortunate inheritance, as well as an almost unique environment in which to realize it.
They also have to thank their mother for their education, and this is perhaps the most surprising aspect of their life. Quite soon after arriving in Botswana, Kate decided to teach them herself. A brave decision, I think I would have counselled against it. But I would have been wrong. Although all their schooling is done in camp, they keep proper terms, have challenging homework assignments and work towards inter- nationally accredited exams. Kate gets good results by standard educational certifications, while at the same time tending, indeed enhancing, the natural sense of wonder that normal children too often lose during their teens. I don't think any reader of these pages could fail to judge her unorthodox School in the Bush a brilliant success. * The proof lies in the book, for, to repeat, the children, and they alone, wrote it. All three authors show themselves to be excellent writers: sensitive, literate, articulate, intelligent and creative.
Kate's choice of Botswana rather than anywhere else in Africa was for- tuitous. It eventually led to her meeting Pieter Kat. And of course the lions - wild lions, living and dying in the world for which the natural selection of their ancestors had prepared them. Pieter is the ideal step-
? Further testimony to the success of Kate's bush school is that Travers and Angus have both been accepted by the (first-class) universities of their choice, UC Santa Barbara and Stanford, respectively.
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? father for her children, and these young scientists have in turn become an indispensable part of the lion research and conservation project.
It wasn't till last year that my family and I finally visited the camp. The experience was unforgettable, and I can testify to the picture painted in The Lion Children. It really is just like that: more wonderful than mad, but a bit of both. My daughter Juliet went out ahead, part of a large invasion of young visitors who soon picked up the enthusiasm of the resident family. On Juliet's first full day in Africa, Travers took her out in a Land Rover, tracking radio-collared lions. When we received Juliet's letter home, brimming with excitement at such an initiation, I relayed the story to her grandmother, who interrupted me with panic in her voice: 'Plus, of course, at least two armed African rangers?
