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SCIENCE-FOR-THE-PEOPLE  June 2002

SCIENCE-FOR-THE-PEOPLE June 2002

Subject:

More on Gould

From:

NEWMAN STUART <[log in to unmask]>

Reply-To:

Science for the People Discussion List <[log in to unmask]>

Date:

Fri, 14 Jun 2002 13:49:03 -0400

Content-Type:

text/plain

Parts/Attachments:

Parts/Attachments

text/plain (148 lines)

                        To appear in Journal of Biosciences vol. 27(4),
September 2002

Stephen Jay Gould
by
Stuart A. Newman
Department of Cell Biology and Anatomy
New York Medical College
Valhalla, NY 10595 USA
(Email, [log in to unmask] <mailto:[log in to unmask]>)

On May 20th the world lost a major thinker in the field of evolutionary
biology and its foremost historian of ideas in this field.  Stephen Jay
Gould's prodigiousness, breadth of knowledge and grandiloquence sometimes
made him seem more like a figure of the 19th  century than the 20th, but his
work was critical in the transition from the neo-Darwinian synthesis to the
more developmentally-influenced concepts of evolution of the present period.


Gould was a keen observer of natural phenomena, and fearless in pursuing the
consequences of his observations even if they led to unorthodox conclusions.
Notwithstanding his many talents, he did not involve himself in the
molecular approaches that emerged as the mainstream of evolutionary and
developmental studies during his lifetime.  This worked to his advantage,
however, in that it freed him to consider large scale macroevolutionary
events through the telescope of paleontology, to which he brought a new
glamour, rather than the microscope of molecular genetics, where the big
picture is frequently missed.  Punctuated equilibrium-the fits and starts of
the fossil record-was elevated by Gould and his colleague Niles Eldridge to
a phenomenon that required rethinking the neo-Darwinian orthodoxy (Gould and
Eldredge, 1977). So was the "Burgess Shale effect," the finding that most,
if not all, the major animal body plans burst forth in a relatively narrow
span of time between 500-600 million years ago (Gould, 1989).

The standard neo-Darwinian view (deriving primarily from theoretical work of
R.A. Fisher) was that sudden morphological change (saltation) cannot be
characteristic of evolution and there must thus be something incomplete
about observations of morphological discontinuity-a consequence of gaps in
the fossil record, and so forth. Most viable genetic changes in modern
organisms are of small effect, leading to incremental phenotypic change.
Mutations of large effect, seemingly needed for both punctuated equilibrium
and the Burgess Shale phenomenon, are typically pathological and so, Fisher
argued, won't be fixed (established) in natural populations.

While neo-Darwinism had ways of explaining the emergence of genetically
distinct populations, such as geographic isolation (Ernst Mayr) and genetic
drift on multipeak adaptive landscapes (Sewell Wright), and therefore could
accommodate the "punctuated" character of the fossil record, all such models
had more difficulty with the "equilibrium" aspect.  Why is morphology so
static in the face of large-scale external changes?  Gould and Eldredge's
insight concerning the tempo and mode of evolution pointed to the need for
different kinds of answers.

Gould typically used metaphors rather than mechanistic analyses to argue his
case.  The spandrels of San Marco, the entry point into a famous critique of
adaptationism by Gould and his colleague Richard Lewontin (Gould and
Lewontin, 1979), are structural side effects of architectural necessities
that later can be used for other purposes.  This phenomenon in the evolution
of organisms was later generalized as the concept of  "exaptation" by Gould
and Elisabeth Vrba (Gould and Vrba, 1982).  Another metaphor Gould was fond
of was "Galton's polyhedron" (Gould, 1986).  A sphere will roll along a
surface by making incremental changes in its position of contact. A
polyhedron, however, if prodded, will make a quantal change in its contact
with the surface.  By such didactic means Gould sought to establish that
there were alternatives to incremental Darwinism.

In search of predecessors in his approach to macroevolution, and with
characteristic intrepidness, Gould revisited the work of such unjustly
scorned figures in biology as Jean-Baptiste Lamarck, the first scientific
thinker about organic evolution (Gould, 1979; 1999a,b), D'Arcy W. Thompson,
who believed that biological forms bore the imprint of physical forces
(Gould, 1971), and Richard Goldschmidt, coiner of the term "hopeful
monster," a shorthand for certain saltationist views now coming back into
fashion (Gould, 1977a).  But the fact that his own scientific specialty
afforded him no causal means for accounting for the macroevolutionary
phenomena that drew his attention made him a ready target for neo-Darwinist
critics.  His remoteness from the contemporary cellular and material
sciences that might yield such explanations also kept him from attaining the
status of revolutionizing evolutionary theorist to which, from the evidence
of his writings, particularly his last book (Gould, 2002), he clearly
aspired.

Gould's most serious attempt to deal with the actual underlying basis of
macroevolution was in his first book "Ontogeny and Phylogeny" (Gould, 1977b)
where he drew on 19th and early 20th century analyses of comparative anatomy
and embryology to show that subtle changes in timing of developmental
process can lead to morphological novelty through a process termed
"heterochrony." This work was neglected by the mainstream of developmental
biology, which was then embarking on a productive but narrow fascination
with genetic mechanisms of development. But Gould's view that developmental
mechanics (not just genetics) was important for understanding morphological
evolution was taken seriously by some working at the interface of
evolutionary and developmental biology, and the book had a surreptitious
career as an important spur to the new field of "evo-devo" that has emerged
over the last decade. This field also has its share of what Gould termed
"Darwinian fundamentalists" (Gould, 1997) but that style of thought is
losing force as extensive discordances between genetic and morphological
evolution are coming to light, and nongenetic determinants of phenotypic
evolution (physical properties of tissues, emergent properties of complex
systems) are entering into the mix.

Stephen Gould's intellectual audacity was undoubtedly tied to his political
progressivism.  An avowed Marxist, he assumed the role of public citizen in
opposing the misuse of biology in social thought and policy (Gould, 1981)
and the corruption of science education promoted by religiously motivated
creationists (Gould, 1999c).   As with his scientific work, his public role
was informed by an historical appreciation of the origins of received
doctrines, along with a unique capacity to see what they left unexplained
and to imagine better possibilities.

References

Gould S J 1971 D'Arcy Thompson and the science of form; New Literary History
II 2
        229-58
_______ 1977a Ontogeny and Phylogeny  (Cambridge MA: Harvard University
Press)
_______ 1977b The return of hopeful monsters; Natural History, 86
(June-July): 22-30
_______ 1979 Shades of Lamarck;  Natural History 88 (August): 22-28
_______ 1981 The Mismeasure of Man  (New York: W. W. Norton)
_______ 1986 The egg-a-day barrier; Natural History 95 (July) 16-24
_______ 1989 Wonderful Life (New York: WW Norton)
_______ 1997 Darwinian fundamentalism; New York Review of Books (June 12)
34-37
_______ 1999a A division of worms; Natural History 108 (February) 18-22;
76-81
_______ 1999b Branching through a wormhole; Natural History 108 (March)
24-26; 84-
                89
_______ 1999c Dorothy, it's really Oz. A pro-creationist decision in Kansas
is more
than a blow against Darwin  Time Magazine (August 23) 59
_______ 2002 The Structure of Evolutionary Theory (Cambridge MA: Harvard
Univ.
Press)
_______ and Eldredge N  1977 Punctuated equilibria: The tempo and mode of
evolution
reconsidered; Paleobiology 3 115-51
_______ and Lewontin R C 1979 The spandrels of San Marco and the Panglossian

paradigm: a critique of the adaptationist programme; Proc. R. Soc. Lond. B
205
581-98
_______ and Vrba E 1982 Exaptation-a missing term in the science of form;
Paleobiology 8 4-15

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