When You Should Suspect Selection: When Adaptive Complexity Exists
"Adaptive complexity" describes any system composed of many interacting parts where the details of the parts' structure and arrangement suggest design to fulfill some function. The vertebrate eye is the classic example. The eye has a transparent refracting outer cover, a variable-focus lens, a light-sensitive layer of neural tissue lying at the focal plane of the lens, a diaphragm whose diameter changes with illumination level, muscles that move it in precise conjunction and convergence with those of the other eye, and elaborate neural circuits that respond to patterns defining edges, colors, motion, and stereoscopic disparity. It is impossible to make sense of the structure of the eye without noting that it appears as if it was designed for the purpose of seeing -- if for no other reason that the man-made tool for image formation, the camera, displays an uncanny resemblance to the eye. Before Darwin, theologians, notably William Paley, pointed to its exquisite design as evidence for the existence of a divine designer. Darwin showed how such "organs of extreme perfection and complication" could arise from the purely physical process of natural selection.
The essential point is that no physical process other than natural selection can explain the evolution of an organ like the eye. The reason for this is that structures that can do what the eye does are extremely low-probability arrangements of matter. By an unimaginably large margin, most objects defined by the space of biologically possible arrangements of matter cannot bring an image into focus, modulate the amount of incoming light, respond to the presence of edges and depth boundaries, and so on. The odds that genetic drift, say, would result in the fixation within a population of just those genes that would give rise to such an object are infinitesimally small, and such an event would be virtually a miracle. This is also true of the other nonselectionist mechanisms outlined by Gould and Lewontin. It is absurdly improbable that some general law of growth and form could give rise to a functioning vertebrate eye as a by-product of some other trend such as an increase in size of some other part. Likewise, one need not consider the possibility that some organ that arose as an adaptation to some other task, or a spandrel defined by other body parts, just happened to have a transparent lens surrounded by a movable diaphragm in front of a light-sensitive layer of tissue lying at its focal plane. Natural selection -- the retention across generations of whatever small, random modifications yield improvements in vision that increase chances of survival and reproduction -- is the only physical process capable of creating a functioning eye, because it is the only physical process in which the criterion of being good at seeing can play a causal role. As such it is the only process that can lead organisms along the path in the astronomically vast space of possible bodies leading from a body with no eye to a body with a functioning eye.
This argument is obviously incomplete, as it relies on the somewhat intuitive notion of "function" and "design." A skeptic might accuse the proponent of circularity, asking why a lump of clay should not be considered well-designed to fulfill the function of taking up exactly the region of space that it in fact takes up. But the circle can be broken in at least three ways. First, biologists need posit far fewer functions than there are biological systems; new functions are not invented for each organ of each organism. Furthermore, each legitimate function can be related via a direct plausible causal chain to other functions and -- critically -- to the overall function of survival and reproduction. Finally, convergent evolution and resemblance to human artifacts fulfilling the same putative function give independent criteria for design. But regardless of the precise formulation of the modern argument from design (see, e.g., Cummins, 1984), it is not controversial in practice. Gould himself readily admits that natural selection is the cause of structures such as the vertebrate eye, and he invokes the criterion of engineering design, for example, to rescue Darwinism itself from the charge of circularity (Gould, 1977a). Presumably this is why Gould and Lewontin concede that they agree with Darwin that natural selection is "the most important of evolutionary mechanisms."
Cummins, R. (1984) Functional analysis. In: Conceptual issues in evolutionary biology, ed., E. Sober. MIT Press.
Gould, S. J. (1977a) Darwin's untimely burial. In: Ever since Darwin: Reflections on natural history, ed., S. J. Gould. Norton.
from "Natural Language and Natural Selection"
Quoted on Sat Sep 28th, 2013