The controversial fruits of a mind full of spandrels

Reminiscent of the late English playwright Dennis Potter's convulsive exit from this life with two final television plays in …

Reminiscent of the late English playwright Dennis Potter's convulsive exit from this life with two final television plays in 1994, the great American essayist and Harvard palaeontologist, Stephen Jay Gould, who died earlier this year from cancer, wrote his own epitaph with these two books: his final compendium of essays (many taken from his monthly slot in Natural History); and his magnum opus, The Structure of Evolutionary Theory, a deeply serious door-stopper indeed.

Both books cap an extraordinary career as an evolutionary theorist and writer who, over 30 years, produced more than 20 books and nearly 1,000 scientific papers. The title of his last collection, I Have Landed, is taken from an inscription his maternal grandpaw wrote in a grammar book when the latter arrived as a child immigrant, along with his extended Hungarian Jewish family, at Ellis Island on September 11th, 1901. (Apparently, Gould was flying from Milan to NYC on September 11th, 2001, to commemorate this centenary with his mother, before the Twin Towers atrocity landed him in Halifax).

Typically, these essays centre on evolution, with Gould's trademark diversions into art, architecture, literature, history and baseball. There are characteristic gems: the piece on novelist and lepidopterist Vladimir Nabokov who, like Gould, worked at Harvard's Museum of Comparative Zoology and found the "precision of poetry in taxonomic description". In like mode, Gould muses on nature as conceptualised through art and science: setting the sublime landscape paintings of Frederic Edwin Church against the writings of Alexander von Humboldt, inventor, navigator, cartographer and naturalist.

Elsewhere, Gould scolds naturalist Ernst Haeckel (1834-1919) for shoehorning evidence into wrong-headed theories, not least Haeckel's eugenical assertions of German cultural and biological superiority - with Gould, as ever, refusing to discuss any application of genetics to human society. Gould is less shy in roasting the Kansas Board of Education for removing, in 1999, evolution (and Big Bang theory) from the state science curriculum. Gould evilly quotes Dorothy in The Wizard of Oz: "They still call it Kansas, but I don't think we're in the real world any more . . ."

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But if Gould's "popular" writing style became increasingly prolix in later years, try grappling with The Structure of Evolutionary Theory, a megatome whose title, Gould admits, is partly a reference to European "structuralist" thinkers, rather than the "pan-adaptationist" British school.

Unfortunately, this book is not a generalised summation of the state of knowledge of the history of life on Earth - nor even of how evolution is theoretically framed - but rather a jargonised restatement of Gould's controversial, often narrow positions within evolutionary theory. Echoing his idol, Charles Darwin, he calls it "one long argument", where he ploughs through mountains of selective written sources.

Gould starts from Darwin's statement in a letter to naturalist Hugh Falconer that, although subsequent discoveries might prove Darwin wrong in detail, the framework of his theory would last. Falconer replied that Darwin had built the foundation of a great edifice, comparing it to Milan's Duomo, begun in 14th-century Gothic style, and finished in a kind of glued-on, late 16th- century Baroque style.

This pedantic-semantic distinction sets the tone for Gould's own rather baroque edifice, which, although shoring up Darwin, discounts the latter's assertion that the slow, "gradualist" view of evolution is sufficient - even over vast, geological time - to account for the staggering diversity of life on Earth. Instead, Gould insists on a whole other macroevolutionary tier of "patterning forces" - which he rarely identifies.

Gould sketches the development of these arguments from even before Darwin's On the Origin of Species (1859) - from geologist Charles Lyell's "uniformalist" theory; through the rediscovery of Mendelian heredity around the turn of the 20th century; to what he calls the "hardening of the Modern Synthesis".

Gould devotes nearly 300 pages to the assault he made on this supposed orthodoxy in a 1972 paper, written with Niles Eldredge. Their idea of "punctuated equilibrium" re-read the fossil record in terms of long periods of evolutionary "stasis", interrupted by periods in which many radically new species emerged, perhaps sparked by catastrophic environmental change caused by, say, asteroidal impact (Gould greedily milks the growing evidence for the bolide that may well have eradicated the dinosaurs).

Critics have long claimed that "punctuated equilibrium" is less an original concept than a tweaking of parameters (one of Gould's "gradualist" detractors called this "evolution by jerks"; Gould famously riposted "evolution by creeps"). But apart from acrimonious debate, Gould maintains that punctuated equilibrium inspired much research into the overwhelming predominance of stasis in the fossil record (often erroneously written up as evidence for gradualism) - even over periods of enormous environmental change.

Gould exhaustively restates his opposition to the "pan-adaptationist" view which, when examining the traits of an organism (from basic morphology to behaviour), primarily looks for adaptive or reproductive advantages. Gould reckoned it was more important to identify developmental mechanisms which imposed structural and historical restraints on evolutionary change.

For many researchers, Gould thus created an artifical opposition between what were essentially complementary factors in the same broader evolutionary process. In his lifetime, this ignited many colourful spats about the adaptive/reproductive merits (or otherwise) of the parenting habits of the blue-footed booby; the sexual suicides of male redback spiders; the erectile pseudopenis, and indeed pseudoscrotum, of the female spotted hyena - let alone the female human orgasm.

(According to Gould, there is no more need to explain female orgasm in adaptive terms than there is to explain male nipples, both being by-products of developmental processes. Opponents batted back findings that during orgasm, the hungry mouth of the cervix makes spasmodic little lunges - but let's not go into that just now.)

Many traits in nature, declares Gould, are decidedly non-adaptive consequences of evolutionary history. Throughout, he employs the term, "spandrel", which he coined with geneticist Richard Lewontin in 1977. This refers, by metaphorical extension to architectural engineering, to the four "left-over" triangles between the four arches supporting the dome in the San Marco cathedral in Venice - mere by-products of the structural arches, despite being opportunistically illustrated with mosaics of the four evangelists.

Similarly, Gould's logic goes, such "spandrel" by-products of biological architecture, honed by evolutionary time, have often later been co-opted for other purposes in a process he called "exaptation". Classic spandrels include the many outlandish facilities of the human brain. Could natural selection really have anticipated such cultural struggles as writing, never mind having to wade through Gould's Structure of Evolutionary Theory?

Taking a last, affectionate cudgel to his old slugging partner, Richard Dawkins - and the latter's "selfish gene" as the ultimate site of natural selection, with organisms acting merely as blind "vehicles" - Gould argues, fairly sensibly, that natural selection operates at a number of heirachical levels: from genes and their expression and regulation through "cell lineages", individual organisms and demes (local populations of interbreeding organisms) to entire species or clades (groups of species descended from common ancestors).

As though aching for the abstractions of modern physics, Gould's gobfuls of theory often leave one light-headed - even when he explains the hands-on fossil work of sympathetic researchers. Rather frightfully, he blows his own tuba loudly about the international controversy engendered by punctuated equilibrium for a good 50 pages, in which he primly accuses his detractors of jealousy, base emotion and nastiness. And before acknowledging his debt to Thomas Kuhn's The Structure of Scientific Revolutions (1962), he fanfares the prevalence of his idea in literature, histories of civilisation and technology, models of human learning, even political and economic theory.

Many scientists still assert that Gould continually misinformed the public about contemporary evolutionary research, particularly in sub-disciplines which explore behaviour. His anti-adaptationist stance particularly targeted the "sociobiology" ideas of ant-biologist Edward O. Wilson - and one can easily imagine Gould spinning in his grave at the dafter, ongoing vogues within "evolutionary psychology".

Generally, Gould rarely communicates any real sense of biological mechanisms, basic genetics, let alone the cascade of recent discoveries in molecular, developmental biology. In his lifetime, he was often derided for doing little empirical work beyond the analysis of small-scale evolution within populations of West Indian land snails - which, to be fair, he does acknowledge.

Interestingly, Gould characterises evolutionary theory as a "historical science", and in summing up, makes much play of the role of "contingency", a factor so crucial to historians and literary novelists. However, his many references to a philosophy of science are often contradictory, and his repeated claim that science is "culturally embedded" again echoes Kuhn, whose work (like Gould's own) is more often cited in the humanities than in science.

ESSENTIALLY, Gould's big book is part philosophy, part history of ideas, part exegesis of Darwin, part "intellectual autobiography" and part over-blown, rambling speculation. The word "spandrel", for example - particularly when one first encounters it - is an elegant encapsulation of a concept. Yet like much of Gould's terminology, he over-invests it with monolithic import. Worse, the circularity of his metaphors promiscuously span across literary and Old Testament references, architecture, biomorphology and the shaky gantry of evolutionary theory itself - hardly endearing him, I suspect, to most working biologists.

There are uplifting passages in this book - such as Gould's succinct demolition of media mishandlings of the origins-of-man debate. However, his openly partisan outbursts and lengthy digressions - for all their technical verbiage and unwieldy sentences - scarcely conceal the fact that this material is often very woolly, conceptually slight, and repetitive to the point of quasi-dementia. One wonders whether his Harvard editors had any mandate to truly discipline this book into publication.

But let me not tramp the dirt down on Gould as, like many readers, I have relished many a puff of epiphany down the years from his dissections of lived-in concepts and popular misconceptions - let alone the scarifying cheerfulness with which he wrote about his fatal abdominal mesothelioma.

"When I am writing," he wrote, "I feel no pain". In the self-conscious finality of his impending death - like Potter, as I say - he seemed to mystically fuse with his public persona; becoming immaculately subsumed into his world of hazed-out theory and, ultimately, this final text.

I Have Landed: Splashes and Reflections in Natural History. By Stephen Jay Gould. Jonathan Cape, 418 pp. £17.99 sterling

The Structure of Evolutionary Theory. By Stephen Jay Gould. Belknap/Harvard University Press, 1433 pp. £27.50 sterling