Specific defense protects against some parasite genotypes but not others, w
hereas non-specific defense is effective against all genotypes of a parasit
e. Some empirical studies observe hosts with variability only in non-specif
ic defense, other studies find only specific defense. I analyse a model wit
h combined specific and non-specific defense to determine the conditions th
at favor detectable variation in each form of defense. High variation in no
n-specific defense is often maintained when resistance increases in an acce
lerating way with investment, whereas low variation tends to occur when res
istance increases at a decelerating rate with investment. Variation in spec
ific defense rises as the parasite pays a higher cost to attack a broad hos
t range (high cost of virulence), as the number of alternative specificitie
s declines, and as the average level of non-specific defense increases. The
last condition occurs because greater non-specific protection tends to sta
bilize the gene frequency dynamics of specific defense. Selection favors a
negative association between costly components of specific and non-specific
defense-hosts defended by one component are favored if they have reduced a
llocation to other costly components. A negative association confounds the
measurement of costs of resistance. Individuals with specific defense may h
ave reduced investment in costly nonspecific defense. This leads to an appa
rent advantage of specifically defended hosts in the absence of parasites a
nd a measured cost of resistance that is negative. (C) 2000 Academic Press.