The Structure of Evolutionary Theory

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The Structure of Evolutionary Theory Page 153

by Stephen Jay Gould


  For example, Gersick emphasizes the need for active resistance towards change as a validation of stasis, and she makes the same link that Eldredge and I have stressed for biological evolution between punctuational change in hierarchical systems and structural constraint viewed as partly contrary to an [Page 959] adaptationist paradigm. She writes (1991, p. 12): “Gradualist paradigms im­ply that systems can 'accept' virtually any change, any time, as long as it is small enough; big changes result from the insensible accumulation of small ones. In contrast, punctuated equilibrium suggests that, for most of systems' histories, there are limits beyond which change is actively prevented, rather than always potential but merely suppressed because no adaptive advantage would accrue.”

  Gersick's lists of commonalities among her six levels, both for periods of stasis and for episodes of punctuation, satisfy the strategy of conjoints, while her ranked list of scales, and especially her linkages of particular categories to the two most overarching theories of punctuational change — Kuhn's for hu­man thought and Prigogine's for the natural world — meet the criterion of generalization. For example, her chart of comparison among the six levels for “equilibrium periods” cites both commonalities and conjoints, with an inter­estingly different emphasis (from our concerns with biological systems) upon the potential for strong limitation placed upon incremental pathways within a plateau — an important theme for the Lamarckian character of human cul­tural change. She lists as “commonalities” (p. 17): “During equilibrium peri­ods, systems maintain and carry out the choices of their deep structure. Sys­tems make adjustments that preserve the deep structure against internal and external perturbations, and move incrementally along paths built into the deep structure. Pursuit of stable deep structure choices may result in behavior that is turbulent on the surface.”

  Similarly, her chart for punctuational episodes stresses the unpredictability and potential nonprogressionism of outcomes, a surprising theme for hu­man systems based on supposed and explicit goals, but a notion that we did not emphasize in formulating punctuated equilibrium because biologi­cal evolution proceeds in a highly contingent manner for so many other rea­sons (see Gould, 1989c), some recognized and emphasized by Darwin him­self. Thus, this important theme, while equally central within the structure of our theory, did not have similar salience for us — and we thank Gersick for her insight and generalization. Gersick writes in her heading (p. 20): “Revo­lutions are relatively brief periods when a system's deep structure comes apart, leaving it in disarray until the period ends, with the 'choices' around which a new deep structure forms. Revolutionary outcomes, based on inter­actions of systems' historical resources with current events, are not predict­able; they may or may not leave a system better off. Revolutions vary in mag­nitude.”

  As another example of fruitful borrowing across disciplines, Mokyr (1990, p. 350) begins his study of technological change by noting that Alfred Mar­shall's advocacy of gradualism in economics played a similar role in the hu­man sciences to Darwin's impact upon the natural sciences: “Charles Darwin and Alfred Marshall were both extremely influential men . . . Darwin and Marshall both believed that nature does not make leaps. Both were influ­enced by a long and venerable tradition that harked back to Leibniz rooted in the Aristotelian notion of the continuity of space and time.” (On the same [Page 960] subject, see also Loch, 1999, on “A Punctuated equilibrium model of technol­ogy diffusion.”)

  The development of improved systems for human communication in the past century must represent one of the most goal-directed and clearly pro­gressionist sequences identifiable in either the human or natural sciences — hence the apparent consonance with gradualist models, and the unpromising character, at first glance, of punctuationalist alternatives. But Mokyr defends a punctuational reformulation by centering his descriptions and explanations on a comparison of stages in this technological trend with the discrete origin of biological species as genuine entities, and then citing punctuated equi­librium for emphasizing this theme as a reform within Darwinian biology (Mokyr, 1990, p. 351).

  While not denying the clearly goal-oriented and progressive nature of the trend — and while also (along with Gersick) noting the Lamarckian capacity of human culture to change directionally and incrementally within plateaus, but also stressing the qualitative differences between this limited gradualism and the much larger and quicker transitions of goal-directed punctuations — Mokyr (p. 354) describes the basic outline of this history as four stages sepa­rated by punctuational breaks. He also, again as with Gersick, stresses the nonpredictability of outcomes, and then notes, continuing the analogy with speciation, that punctuational models enjoin the study of particular condi­tions favorable to leaps of change, an issue that does not arise in explanations based upon gradualistic anagenesis:

  Long-distance communications thus illustrate the abruptness of technological change. There was no natural transition from semaphore to the electrical telegraph, or a gradual movement from the telegraph to the first radio transmission by Marconi in 1894, nor a smooth natural development from the long-wave radio used in the first thirty years to the shortwave systems of the later 1920s. Each of the three systems was subsequently perfected by a long sequence of microinventions, but these would not have occurred without the initial breakthroughs. Their con­cept was novel, they made things possible that were previously impossi­ble, and they were pregnant of more to come. Therein lies the essence of a macroinvention.... Many macroinventions, just like the emergence of species, were the result of chance discoveries, luck, and inspiration. Biologists agree that certain environments are more conducive to speciation than others.

  Just as for technological change, we tend to view the history of scientific ideas on particular subjects as, in principle, the most incrementally progres­sionist of all human activities by the empiricist paradigm of ever-closer ap­proximation to natural truth through objective accumulation of data under unchanging principles of “the scientific method” — an idea famously chal­lenged by Kuhn (1962) in the most influential punctuationalist theory in 20th century scholarship — see p. 967. Thus, punctuationalist reformulations in this domain tend to strike people as especially surprising. The distinguished [Page 961] Dutch petrologist E. Den Tex (1990), in summing up his career of study on the nature of granites, credited punctuated equilibrium with reorganizing his lifelong attempt to make sense of a complicated history, stretching back to the late 18th century (the neptunist-plutonist debate between the schools of Werner and Hutton), and featuring fluctuations between poles of two dichot­omies (sedimentary vs. igneous formation, and recruitment from deep mag­mas vs. transformation from existing crustal rocks), complicated by shifting allegiances and amalgamations, followed by breakages, of separable aspects of all end-member theories.

  In his fascinating and highly personal paper, entitled “Punctuated equi­libria between rival concepts of granite genesis,” Den Tex (1990) notes that he had first tried to apply other models of noncontinuous and progres­sive change, especially the celebrated Hegelian notion of successive syntheses reached by opposition between a thesis and its antithesis. He then found a better general description, with new and fruitful hints for explanation, in our model of punctuated equilibrium, particularly in the parsing of history as dis­crete steps, analogous to individuated species with definite sets of proper­ties — a process “in dynamic equilibrium ... punctuated from time to time by allopatric speciation, i.e. by rapid, random, discrete steps taking place in lo­cations isolated from the main stem” (1990, p. 216).

  Finally, since punctuated equilibrium arose as a theory about change in the natural world (not in the history of human understanding thereof), Eldredge and I have been gratified by the utility of our theory in suggesting structurally homologous modes of change in other branches of natural science. I have been especially pleased by geological examples distant from our own paleontological concerns, because no other field can match anglophonic geology — resulting largely f
rom the legacy of Lyellian uniformitarianism (see Chapter 6) — in explicit, and often exclusive, fealty to strictly gradualistic models.

  Lawless (1998, and a good name for iconoclasts), in an article entitled “Punctuated equilibrium and paleohydrology,” notes the hold that gradual­istic models have imposed on the history of ideas about hydrothermal ore de­posits, particularly of gold. He begins by expressing a paradox: if ores accu­mulate gradually in such systems, and given the average amount of gold carried in most percolating waters, minable deposits should be much more common — indeed almost ubiquitous in hydrothermal systems that persist for at least 25,000 years. But the much rarer distribution of such deposits suggests to Lawless that periods of accumulation must be limited to brief episodes “which cause vigorous boiling through a restricted volume of the reservoir” (p. 165). Lawless views the general history of hydrothermal sys­tems — including the development of ore deposits as just one feature among many — as punctuational, and caused by rapid, intense, rarely-acting forces: “Such disturbances caused by tectonic activity, magmatism, volcanic activity, erosion, climatic changes or other processes may occur at long intervals, but be responsible for producing some of the most significant characteristics of the system, including economic mineral deposits.”

  As a matter of potential practical importance, Lawless recognizes that, just [Page 962] as punctuated equilibrium must not be construed as an argument against predictable trends, but rather as a different mechanism for the episodic produc­tion of such trends, so too might the punctuational origin of hydrothermal ore deposits be reconceptualized as directional but episodic. He writes (p. 168): “Recognition of the quasi-cyclic and episodic nature of these events within the lifetime of a hydrothermal system could be described as leading to more 'catastrophic' models. There are similarities to the concepts of punctu­ated equilibrium recently proposed in paleontology and biological evolution . . . these concepts emphasize the importance of specific events which are of random occurrence on a short time scale but statistically predictable on a longer scale.”

  TWO CONCLUDING EXAMPLES, A GENERAL STATEMENT, AND A CODA.

  As final examples in this chapter, two recent authors have used punctuated equilibrium as the central organizing principle for books on subjects of differ­ent scale, but of great importance in human life and history — Kilgour (1998) on The Evolution of the Book, and Thurow (1996) on The Future of Capi­talism. Moreover, each author uses punctuated equilibrium not as a vague metaphor, but as a specific model of episodic change offering casual insights through the identification of structural homologies as defined in this chapter.

  Kilgour notes that a theme of greater efficiency marks the history of book making (and might be misread as evidence for anagenetic gradualism, just as trends in the evolutionary history of clades have often been similarly miscon­strued when a punctuational model of successive plateaus defined by discrete events of branching actually applies). He writes (p. 4): “Form aside, the ma­jor change throughout the entire history of the book has been in the continu­ous increase in speed of production: from the days required to handwrite a single copy, to the minutes to machine-print thousands of copies, to the sec­onds to compose and display text on an electronic screen.”

  But, as Kilgour knows, and adopts as the major theme of his book, form cannot be put “aside.” When one probes through these progressive improve­ments in function to underlying bases in form, the history of the book be­comes strongly punctuational. In a pictorial summary for his central thesis (Fig. 9-36), Kilgour views the evolution of the book — defined (p. 3) as “a storehouse of human knowledge intended for dissemination in the form of an artifact that is portable, or at least transportable, and that contains arrange­ments of signs that convey information” — as a sequence of four great punctu­ations: the clay tablet, the papyrus roll, the codex (modern book), and the electronic “book” (with no canonical form as yet since we are now enjoying, or fretting our way through, the rare privilege of living within a punctuation), with three “subspeciational” punctuations within the long domination of the codex (Gutenberg's invention of printing with movable type in the mid 15th century, and the enormous additional increases in production made possible first by the introduction of steam power at the beginning of the 19th century, and then by the development of offset printing in the mid 20th century).

  As I have emphasized throughout this discussion of human cultural and [Page 963] technological change, the Lamarckian nature of inheritance for these pro­cesses permits more directional accumulation within periods of overall stasis in basic design than analogous chronologies of biological evolution can prob­ably exhibit. Thus, while the codex (that is, the familiar “bound book”) en­joyed its millennium and a half of domination in fundamentally unchanged form, several innovations both in design (the introduction of pagination and indexes) and in human practice and collateral discovery (the invention of eye­glasses at the end of the 13th century, and the spread of the “newfangled” practice of silent reading in the 15th century) greatly expanded the utility of a product that remained stable in form. (While books remained scarce, people read aloud and, apparently, did not even imagine a possibility that seems ob­vious to us — namely, that one might read without speaking the words. By reading aloud, one copy could be shared with many, but silent reading de­mands a copy for each participant.)

  But the four great designs (or at least the three we know, for the fourth has not yet stabilized) have experienced histories strikingly akin, in more than vaguely metaphorical ways, to the origin and persistence of biological species treated as discrete individuals. In the first of three major similarities, each principal form persisted in effectively unchanged design for long periods of time by standards of human technological innovation. Moreover, each transi­tion introduced a great improvement by solving an inherent structural prob­lem in the previous design. Therefore, the extended persistence of each flawed

  9-36. Kilgour, in his 1998 work on the evolution of the book, presents a punctuational model of day tablet, to papyrus roll, to codex, to printing, and onward to the undetermined future of the electronic book. All major features of the biological model apply here in adequate isomorphism for causal insight, including the survival of ancestors after the branching of descendants.

  [Page 964]

  design illustrates an important reason, more “environmental” than struc­tural, for the existence of stasis in natural systems: the advantages of incum­bency. “The extinction of clay tablets,” Kilgour writes (pp. 4-5) “was en­sured by the difficulty of inscribing curvilinear alphabet-like symbols on clay”; while “the need to find information more rapidly than is possible in a papyrus-roll-form book initiated the development of the Greco-Roman co­dex in the second century a.d.” Of this predominant stasis, Kilgour writes (p. 4): “Extremely long periods of stability characterize the first three shapes of the book; clay tablets and papyrus-roll books existed for twenty-five hun­dred years, and the codex for nearly two thousand years. An Egyptian of the twentieth century B.C. would immediately have recognized, could he have seen it, a Greek or Roman papyrus-roll book of the time of Christ; similarly, a Greek or Roman living in the second century a.d. who had become familiar with the then new handwritten codex would have no trouble recognizing our machine-printed book of the twentieth century.”

  Secondly, the successive stages do not specify segments of an anagenetic flow (whatever the punctuational character of each introduction), but rather arise as discrete forms in particular areas — thus following the pattern of branching speciation so vital to the validation of punctuated equilibrium, and also meeting the chief operational criterion for distinguishing punctuated equilibrium from punctuated anagenesis: the survival of ancestral forms after the origin of new species. Kilgour notes (p. 158) “clay tablets and papy­rus-roll books coexisted for two thousand years, much as two biological spe­cies may live together in the same environment.” He also notes, both wryly and a bit ruefully, “that
books on paper and books on electronic screens, will, like clay tablets and papyrus books, coexist for some time, but for decades rather than centuries” (p. 159).

  Third, each form — at least before improved communication of the past two centuries made such localization virtually inconceivable — originated in a par­ticular time and place, and in consonance with features of the immediately surrounding environment, a meaningful analog to the locally adaptive origin of biological species. Kilgour writes (p. 4): “the Sumerians invented writing toward the end of the fourth millennium B.C. and from their ubiquitous clay developed the tablet on which to inscribe it. The Egyptians soon afterward learned of writing from the Mesopotamians and used the papyrus plant, which existed only in Egypt, to develop the papyrus roll on which to write.”

  I know Lester C. Thurow as a colleague from another institution in Cambridge, MA, but I had never discussed punctuated equilibrium with him, and was surprised when he used our theory as one of two defining metaphors in his book, The Future of Capitalism. In distinction to most examples in this chapter, Thurow does invoke punctuated equilibrium in a frankly metaphori­cal and imagistic manner, but he also shows a keen appreciation for the con-joints of punctuated equilibrium applied to his subject of macroeconomics. Thurow writes in the context of the collapse of communism in the Soviet Union, and in the survival of capitalism as a distinctive, effectively universal, and perhaps uniquely workable system of human economic organization [Page 965] within a highly technological matrix. Yet capitalism now faces a crisis of reorganization in a world where major change (both natural and social) occurs by punctuated equilibrium and not by slow incrementation. Thurow therefore presents two metaphors from the natural sciences to ground his argument: “To understand the dynamics of this new economic world, it is useful to bor­row two concepts from the physical sciences — plate tectonics from geology and punctuated equilibrium from biology” (p. 6).

 

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