Abstract
STEPHEN JAY Gould and Richard Lewontin introduced the spandrel as a biological metaphor in a 1979 paper entitled “The spandrels of San Marco and the Panglossian Paradigm: a Critique of the Adaptationist Programme”. Architecturally, the curved triangular spaces, called pendentives, formed above and between two neighboring arches in a vaulted roof supported by four arches, are a type of spandrel (Fig. 15.1). Technically, a spandrel is a “space left over” between portions of an architectural structure, such as the vertical spaces between steps on a staircase or the “spandrel courses” on the sides of high-rise buildings, between the windows of one floor and the windows of the next (Fig. 15.2). As Gould explained after he had delved further into the architectural details, many architects use the term only to refer to two-dimensional spaces, while a continental European school does consider three-dimensional forms, like the San Marco pendentives to be spandrels (Gould 1997, 2002, p. 1250).
The river to the ocean goes
A fortune for the undertow
R.E.M.
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Notes
- 1.
When they wrote the “Spandrels” paper, Gould and Lewontin did not use this terminology, and simply applied the general term “spandrel”; this was clarified by civil engineer and architect Robert Mark (1996), in an article entitled “Architecture and Evolution” in The American Scientist. Gould and Lewontin’s terminological vagueness was noted gleefully by Daniel Dennett in his 1995 book Darwin’s Dangerous Idea, which took such a stridently pan-adaptationist view that H. Allen Orr remarked, “Dennett does not so much champion adaptationism as excoriate those biologists who dare question it.” (Orr 1996). See also Ahouse (1998) for a cogent critique of Dennett’s “a priori selectionism”.
- 2.
Voltaire’s Dr. Pangloss, and his idea that this is “the best of all possible worlds” (despite a deluge of events in Candide’s life that prove the opposite) was at least in part a caricature of Leibniz and his philosophy. Voltaire had been deeply moved by the horrific tragedy of the 1755 Lisbon earthquake – after which the terrified residents rushed from their burning and collapsed homes toward the beach, only to be drowned by a tsunami triggered by the earthquake. This event was seminal in turning Voltaire into a bitter opponent of “philosophical optimism”.
- 3.
Voltaire mercilessly mocked the idea of such trade-offs. As Gould and Lewontin remind their readers, Dr. Pangloss told poor Candide not to worry about suffering from a venereal disease. It was “indispensable in this best of worlds. For if Columbus, when visiting the West Indies, had not caught this disease, which poisons the source of generation, which frequently even hinders generation, and is clearly opposed to the great end of Nature, we would have neither chocolate nor cochineal.”
- 4.
The power of steady misinterpretation has not yet relinquished its hold on power: creationists still use Darwin’s pluralistic comments as “evidence” that he repudiated his own theory! One recent example of this is a book by Randall Hedtke entitled “Secrets of the Sixth Edition”, with a photograph of an elderly, white-bearded Darwin gracing the cover… with the photoshopped addition of a finger raised to his lips in a “ssshhhh” gesture. The book is published by a house that specializes in “homeschool resources”.
- 5.
Darwin responds to such arguments with the following – utterly charming – argument: “If beautiful objects had been created solely for man’s gratification, it ought to be shown that before man appeared, there was less beauty on the face of the earth than since he came on the stage. Were the beautiful volute and cone shells of the Eocene epoch, and the gracefully sculptured ammonites of the Secondary period, created that man might ages afterwards admire them in his cabinet?” (Darwin, Origin of Species, p. 253)
- 6.
The tone of the spandrels article rubbed many people the wrong way. Its rhetorical flourishes inspired a virtual cottage industry of criticism, including a collection of essays about the article, entitled Understanding Scientific Prose (University of Wisconsin Press 1993), which dissected the paper from a variety of angles. The tone of some of the discussion of The Spandrels of San Marco, however, has become strangely personal in a way rarely seen in professional scientific writing. In a review of Understanding Scientific Prose (titled “The Scandals of San Marco”, and published in the Quarterly Review of Biology in 1994), Gerald Borgia concluded that “[p]olitical bias remains as the only plausible explanation [sic!!] for Gould’s attacks on adaptation and sociobiology”. Borgia ended his review by declaring “selection the clear winner in the ‘Spandrels’ debate”. As for Gould and Lewontin, Borgia applauded the fact that “fortunately, their elitist attempt to misdirect science failed in the end”. I barely need to point out that these are not arguments.
Spandrels continues to irritate many in the evolutionary biology community nearly four decades after its publication. A 2011 blog post by Jeremy Fox is entitled “Why the Spandrels of San Marco Isn’t a Good Paper”. In his deliberately provocative blog post, Fox argued that the tone of Spandrels is more suited to that of a “deliberately-provocative blog post” (https://oikosjournal.wordpress.com/2011/08/26/why-the-spandrels-of-san-marco-isnt-a-good-paper/, retrieved on January 14, 2016). Fox accused Gould and Lewontin of “cherry-picking” quotes from Darwin in order to bolster their argument; the very clear and deliberate statements by Darwin in this matter, such as those quoted above, show the cherry-picking claim to be utterly spurious. Among other complaints, Fox cited the work of the architectural engineer Robert Marks, who noted that there were various ways known to construct a dome on four arches at the time San Marco was constructed, and the particular method used was the only one stable enough to support such a large dome; the spandrels are, therefore, adaptive. This, of course, does not make for an argument that current utility is aligned with historical origin. Neither does the parody title of David Queller’s article entitled “The Spaniels of St. Marx and the Panglossian Paradox: A Critique of a Rhetorical Programme”, published in the Quarterly Review of Biology in 1995, or the quasi-Gilbert-and-Sullivan song (“I am the very model of a science intellectual” [sic]) included therein. One can dislike someone for being glib, widely read, and at times bombastic. But that doesn’t make him wrong.
- 7.
In The Structure of Evolutionary Theory, Gould pointed out that none other than Friedrich Nietzsche himself emphasized this point in The Genealogy of Morals. Nietzsche argued that it was critically important to separate current use and ethical interpretation of moral constraints in contemporary society from their historical origin (which was, in his view, the will to power). He wrote “that the origin or the emergence of a thing and its ultimate usefulness, its practical application and incorporation into a system of ends, are toto coelo separate; that anything in existence, having somehow come about, is continually interpreted anew, requisitioned anew, transformed and directed to a new purpose” (quoted in Gould 2002, p. 1216). Nietzsche’s argument included biological as well as social structures. “No matter,” he wrote, “how perfectly you have understood the usefulness of any physiological organ (or legal institution, social custom, political usage, art form or religious rite) you have not thereby grasped how it emerged…for people down the ages have believed that the obvious purpose of a thing, its utility, form and shape are its reason for existence: the eye is made to see, the hand to grasp. So people think punishment has evolved for the purpose of punishing” (quoted in Gould 2002, p. 1217).
- 8.
See Irwin et al. (2011) for a recent survey of the evolution of the mammalian lysozyme gene family, including lactalbumin.
- 9.
See, for example, S. Ohno, Evolution by Gene Duplication (1970) for an important early discussion of this topic.
- 10.
Gould and Vrba suggest that the literature of sociobiology would undergo a “constructive collapse”, which would “vastly broaden our range of hypotheses” if it incorporated the idea of exaptations. Certainly it would eliminate “unprovable reveries about primal fratricide on the African savanna or dispatching mammoths at the edge of great ice sheets”, among other just-so stories giving causal evolutionary explanations for aspects of modern human behavior.
- 11.
For more detail on this fascinating topic, see Wistow’s 1995 book Molecular Biology and Evolution of Crystallins: Gene Recruitment and Multifunctional Proteins in the Eye Lens, and Piatigorsky’s 2007 book Gene Sharing and Evolution.
- 12.
It might be more precise to say that it has the potential to be a spandrel, if a feature is only defined as a spandrel once has been coopted to serve an exaptive function.
- 13.
See note 12; the same qualification applies here.
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Bahar, S. (2018). Spandrels, Exaptations, and Raw Material. In: The Essential Tension. The Frontiers Collection. Springer, Dordrecht. https://doi.org/10.1007/978-94-024-1054-9_15
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