Evolution__3rd_Edition

290 PART 3 / Adaptation and Natural Selection
experimental method, see the special issue of American Naturalist, a supplement to
vol. 154 (July 1999).
On constraints, Antonovics & van Tienderen (1991) look at terminology. Barton &
Partridge (2000) look at the topic in general. On “ghost” adaptations like the neotropical
fruit, see the popular book by Barlow (2000). Byers (1997) is an example discussing
the social behavior of the American pronghorn and Macgregor (1991) reviews the
remarkable genetic constraint in the crested newt and refers to earlier work.
On developmental constraint, Maynard Smith et al. (1985) and Gould (2002b) are
major reviews. McKenzie & Batterham (1994) and McKenzie (1996) discuss the insecticide
resistance example (see also the further reading in Chapter 5, p. 135). Antibiotic
resistance in microbes is a related topic. Levin et al. (2000) discuss how compensatory
mutations that reduce the harmful side effects of the initial resistance mutations may
influence the persistence of antibiotic resistance. The arguments are related to those in
Box 5.2 (p. 119). On developmental stability in general, see Lens et al. (2002). Harvey &
Pagel (1991) contains an account of, and references to, recent work on allometry.
Chapter 9 has further references for canalizing selection. Chapter 20 looks at evolutionary
development, which probably provides the concepts for future studies of
developmental constraint. Galis et al. (2001) discuss the special case of constraints
on digit numbers.
Certain human genes confer resistance to disease, but are otherwise disadvantageous.
These genes probably illustrate constraints due to history (they evolved
recently) and to trade-offs (disease resistance is so important that other adaptations are
compromised). Schliekman et al. (2001) give some calculations for three such genes:
CCR5 − (resistance to HIV), hemoglobin S, and ∆32 (resistance to bubonic plague).
On definition, see the references already given to Williams (1966) and Reeve &
Sherman (1993). I have extracted them, along with another good discussion by Grafen,
in Ridley (1997). A further distinction is between historic and non-historic definitions.
Gould has argued that only characters that retain a constant function should be called
adaptations. See Gould (2002b) for a thorough recent statement of his view, and Reeve
& Sherman (1993) for problems with it.
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