The Fruitful Facets of Gallon's Polyhedron 383
orthogenesis (and of formalism in general) in our struggle to develop a more
adequate modern theory of evolution.
World of Pigeons CO. Whitman: An Orthogenetic Dove in Darwin's
WORLD OF PIGEONS
The old benediction—"may you live in interesting times"—has been regarded as
either a blessing or a curse. Charles Otis Whitman certainly merited such an
epithet, for his professional life spanned the greatest range of opposites through the
grandest transition in ideas that biology has ever experienced. He began by
studying with Louis Agassiz, last of the great and legitimate creationists, and
ended as the chief American promoter of mechanistic embryology in the German
tradition. He made his primary reputation in "cell lineage" studies of the fates and
products of the earliest blastomeres. But, unlike most experimentalists of the time,
he also pursued other research as a gifted natural historian and evolutionary
theorist. His major work in later years, while he served as professor at the
University of Chicago, treated a subject that could not have been more canonical
for Darwinian evolutionary biology in the naturalistic tradition—heredity,
variation, and evolution in Darwin's own chosen organism, the domestic pigeon.
Whitman, however, used Darwin's pigeons to support orthogenesis, and to deny
selection a primary or formative role in evolution.
C. O. Whitman died in 1910, of pneumonia contracted after working furiously
on the first cold day of winter to provide shelter for his birds. (F. R. Lillie, once his
assistant and later his successor at Woods Hole, eulogized his old boss: "In his zeal
for his pigeons, he forgot himself.") Whitman had never published an extensive
defense of his orthogenetic theories. His diverse and voluminous writings were
finally collated and published posthumously as a large three-volume monograph by
the Carnegie Institute of Washington in 1919. The evolutionary debates of the
early 20th century had been fierce, and finally won by the followers of Darwinian
theory. I have often wondered how this history might have differed if this
paramount biologist (Kellogg, 1907, p. 288, called Whitman "the Nestor of
American zoologists") had lived to publish what might have been the best
empirical defense of orthogenesis. In any case, the posthumous and much delayed
1919 monograph was too disjointed, too incomplete and above all, too late, to win
any potential influence.
Whitman accepted Kellogg's classification of evolutionary theories as
auxiliary or alternative to Darwinism. He also agreed with Kellogg that three major
alternatives fueled the great debate as the century turned: Lamarckism,
orthogenesis, and macromutationism. Since Whitman rejected the inheritance of
acquired characters with all Weismann's zeal, his own list of viable alternatives
included only two theories. Since he also believed in the strict continuity of
Darwinian gradualism, de Vries's mutationism held no appeal for him either
(though he regarded the theory as a viable contender, while treating Lamarckism as
a dead issue). These rejections left only orthogenesis as a