Evolution__3rd_Edition

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hypothesis of adaptation or constraint, by direct
measures of selection, by seeing whether the character
is variable and whether the variation is heritable and
can be altered by artificial selection, and by examining
comparative trends.
10 The methods of analyzing adaptation are valid
when applied to adaptive characters and interspecific
trends; they might be misleading for non-adaptive
characters and trends.
11 Not all the effects of an organ will have evolved as
Further reading
CHAPTER 10 / Adaptive Explanation 289
adaptations by natural selection. Some will be inevitable
consequences of the laws of physics.
12 Biologists disagree about how exact, and how
widespread, adaptation is in nature.
13 There are criteria to distinguish adaptive from
non-adaptive characters. Measurement of selection
provides an objective criterion, but is not always practical.
The inherent engineering design of a character is
not always an objective crtierion, but is widely applicable.
The two criteria are closely related.
Williams (1966) is a classic work on adaptation. Gould & Lewontin (1979) is an
influential paper that criticizes the way adaptation has often been studied; Cain (1964)
argues the opposite. Pigliucci & Kaplan (2000) look at 20 years of discussion about
Gould & Lewontin (1979). Lewontin (2000) and Gould (2002b) variously update their
viewpoints. Reeve & Sherman (1993) is a stimulating paper about adaptation. Dawkins
(1982, 1986, 1996) argues that only natural selection can explain adaptation; the 1986
and 1996 books are written for a wide audience. Dennett (1995) is also written for a
broad audience and discusses several of the topics covered in this chapter.
Allen et al. (1998) have compiled an anthology of classic papers about adaptation.
My evolution anthology contains a section of extracts about adaptation (Ridley 1997)
and Rose & Lauder (1996) have edited a multiauthor volume on the topic.
The natural theologian’s argument from design was philosophically undermined by
Hume in his Dialogues Concerning Natural Religion, which are in print in various
paperback editions and (unlike some of Hume’s other philosophical writings) readily
intelligible. I include the passage in Ridley (1997). However, Hume’s abstract argument
did not convince people and it was Darwin’s mechanistic theory of natural selection
that historically toppled that long tradition of thought. See Simpson (1944, 1953) on
orthogenesis.
Dawkins (1996) includes a popular account of Nilsson & Pelger’s (1994) paper
about eye evolution. Land & Nilsson (2002) is a book about animal eyes. Nitecki (1990)
is a multiauthor book about evolutionary innovations. On feathers, see Prum & Brush
(2002) and their references. On preadaptation in general, see also the popular essay by
Gould (1977b, chapter 12). Gerhart & Kirschner (1997) discuss the lactose example.
On the genetics of adaptation, Leigh (1987) includes an account of Fisher’s argument.
Travisano (2001) discusses the emerging research program with microbial
experimental systems.
The methods of studying adaptation are discussed (in addition to the multiauthor
volumes referred to above) by Orzack & Sober (1994), Harvey & Pagel (1991), Parker
& Maynard Smith (1990), Maynard Smith (1978), and Rudwick (1964). For the