Gould, The Panda's Peculiar Thumb

Marsh Epipactis, lower sepals removed

Runway of labellum depressed
after insect lands
Runway of labellum raised
after insect crawls into cup below
D. L. Cramer

that the bigger radial sesamoid may have been fashioned by a very simple genetic change, perhaps a single mutation affecting the timing and rate of growth.

In a panda's foot, the counterpart of the radial sesamoid, called the tibial sesamoid, is also enlarged, although not so much as the radial sesamoid. Yet the tibial sesamoid supports no new digit, and its increased size confers no advantage, so far as we know. Davis argues that the coordinated increase of both bones, in response to natural selection upon one alone, probably reflects a simple kind of genetic change. Repeated parts of the body are not fashioned by the action of individual genes--there is no gene "for" your thumb, another for your big toe, or a third for your pinky. Repeated parts are coordinated in development; selection for a change in one element causes a corresponding modification in others. It may be genetically more complex to enlarge a thumb and not to modify a big toe, than to increase both together. (In the first case, a general coordination must be broken, the thumb favored separately, and correlated increase of related structures suppressed. In the second, a single gene may increase the growth rate in a field regulating the development of digits.)

The panda's thumb provides an elegant zoological counterpart to Darwin's orchids. An engineer's best solution is debarred by history. The panda's true thumb is committed to another role, too specialized for a different function to become an opposable, manipulating digit. So the panda must use parts on hand and settle for an enlarged wrist bone and a somewhat clumsy, but quite workable, solution. The radial thumb is, to use Michael Ghiselin's phrase, a contraption, not a lovely contrivance. But it does its job and excites our imagination all the more because it builds on such improbable foundations.

Darwin's orchid book is filled with similar illustrations. The marsh Epipactis, for example, uses its labellum--an enlarged petal--as a trap. The labellum is divided into two parts. One, near the flower's base, forms a large cup filled with nectar--the object of an insect's visit. The other, near the flower's edge, forms a sort of landing stage. An insect alighting on this runway depresses it and thus gains entrance to the nectar cup beyond. It enters the cup, but the runway is so elastic that it instantly springs up, trapping the insect within the nectar cup. The insect must then back out through the only available exit--a path that forces it to brush against the pollen masses. A remarkable machine but all developed from a conventional petal, a part readily available in an orchid's ancestor.

Darwin then shows how the same labellum in other orchids evolves into a series of ingenious devices to ensure cross-fertilization. It may develop a complex fold that forces an insect to detour its proboscis around and past the pollen masses in order to reach nectar. It may contain deep channels or guiding ridges that lead insects to both nectar and pollen. The channels some-times form a tunnel, producing a tubular flower. All these from a part that began as a conventional petal in some ancestral form. Yet nature can do so much with so little that it displays, in Darwin's words, "a prodigality of resources for gaining the very same end, namely, the fertilization of one flower by pollen from another plant."

Darwin's metaphor for organic form reflects his sense of wonder that evolution could fashion such a world of diversity and adequate design with such limited raw material:

Although an organ may not have been originally formed for some special purpose, if it now serves for this end we are justified in saying that it is specially contrived for it. On the same principle, if a man were to make a machine for some special purpose, but were to use old wheels. springs, and pulleys, only slightly altered, the whole machine, with all its parts, might be said to be specially contrived for that purpose. Thus throughout nature almost every part of each living being has probably served, in a slightly modified condition, for diverse purposes, and has acted in the living machinery of many ancient and distinct specific forms.

We may not be flattered by the metaphor of refurbished wheels and pulleys, but consider how well we work. Nature is, in biologist Francois Jacob's words, an excellent tinkerer, not a divine artificer. And who shall sit in judgment between these exemplary skills?

Stephen Jay Gould teaches biology, geology, and the history of science at Harvard University.

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