The Structure of Evolutionary Theory

(Michael S) #1

220 THE STRUCTURE OF EVOLUTIONARY THEORY


change or extension, radically altered his conception into a full selectionist theory
of hierarchy, the first such proposal in the history of evolutionary thought.
Taking an opposite tack from his original formulation, Weismann now
proposed to define the scope of germinal selection by what such a process could
accomplish without and beyond organismal selection: "We shall attempt to gain
clearness as to what it can do, and how far the sphere of its influence extends, and,
in particular, whether it can effect lasting transformations of species without the
cooperation of personal selection, and what kind of variations we may ascribe to it
alone" (1903, vol. 2, p. 126).
Weismann now recognized that he could use germinal selection to escape the
straitjacket of Panglossian adaptation. He could finally admit the non-adaptive
character of some phenotypic features without straying from the Allmacht of
selection, or giving comfort to Lamarckism—for traits arising by germinal
selection may be neutral or even harmful to survival in the Darwinian world of
competition among organisms. Weismann recognized two classes of nonadaptive
features potentially ascribable to germinal selection.
NEUTRAL FEATURES OF SMALL OR NO IMPORTANCE. Since germinal selection
can promote changes invisible to organismal selection, Weismann wondered
whether such minor variations as human racial differences— attributed by Darwin
to sexual selection based on disparate standards of beauty arising for capricious
reasons in various societies—might actually arise as effects of germinal selection:
"It cannot be denied that there are characters which have no special biological
significance [although] ... it is difficult and often impossible to point these out with
certainty. The shape of the human nose and the human ear, the color of the hair and
the iris, may be such indifferent characters whose peculiarities are to be referred
solely to germinal selection" (1903, vol. 2, p. 134).
ORTHOGENETIC DRIVES THAT MAY YIELD HARMFUL FEATURES AND EVEN LEAD
TO EXTINCTION. In a remarkable departure from the almost strident panselectionism
of his earlier years, Weismann now approved certain claims for orthogenesis, and
admitted the existence of harmful trends (based on directionality in variation) that
organismal selection could not reverse. He even accepted the classic examples of
the anti-Darwinian schools as orthogenetic and harmful prima facie—the antlers of
Irish Elks, and the massive canines of saber-toothed cats (1903, vol. 2, p. 139). He
embraced the best cases of his opponents because germinal selection—once he
reversed his original view and grasped its power to work against organismal
selection— could convert these enemy troops to the doctrine of Allmacht. For
when germinal selection acts with sufficient power, then all the determinants
favored by organismal selection may be eliminated entirely, leaving only the
vigorous determinants of harmful orthogenetic features, and rendering
conventional selection impotent for lack of raw material: "In this case the
variation-direction which had gained the mastery in all ids could no longer be
sufficiently held in check by personal selection, because the variations in the
contrary direction would be much too slight to attain to selective value" (1903, vol.
2, p. 139).

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