A further requisite for the evolution of inducible defense systems is heritable
variation in the degree of inducibility (Karban and Myers 1989; Agrawal 1999).
Genetic variation has frequently been observed in natural populations, e.g., for physical
(trichome density) or chemical (glucosinolate content) resistance characters in
Arabidopsis, and both traits are associated with fitness costs (Mauricio 1998). For
induced resistance traits, however, a fitness benefit has been demonstrated in only
a few cases. One example is radish plants that were induced to accumulate higher
levels of glucosinolates and to produce trichomes at increased density. Compared to
control plants, these induced plants exhibited both increased resistance to herbivory
and increased seed mass (a correlate of lifetime fitness). This experiment confirmed
a role in direct defense for trichomes and glucosinolates as inducible physical and
chemical resistance factors, respectively (Agrawal 1998, 1999). Likewise, in Nicotiana
attenuata, the induced production of nicotine as a chemical resistance factor
was associated with metabolic costs, but provided a fitness benefit when plants were
under attack by herbivores (Baldwin 1998; see also Steppuhn and Baldwin this volume).
Although these findings should not be generalized and a defensive role should
not be assumed for all plant responses to wounding and herbivory, the prevalence
of inducible resistance traits in present day plant-herbivore systems implies that
such responses are likely the result of natural selection imposed by insect herbivores
during evolution.
A further requisite for the evolution of inducible defense systems is heritablevariation in the degree of inducibility (Karban and Myers 1989; Agrawal 1999).Genetic variation has frequently been observed in natural populations, e.g., for physical(trichome density) or chemical (glucosinolate content) resistance characters inArabidopsis, and both traits are associated with fitness costs (Mauricio 1998). Forinduced resistance traits, however, a fitness benefit has been demonstrated in onlya few cases. One example is radish plants that were induced to accumulate higherlevels of glucosinolates and to produce trichomes at increased density. Compared tocontrol plants, these induced plants exhibited both increased resistance to herbivoryand increased seed mass (a correlate of lifetime fitness). This experiment confirmeda role in direct defense for trichomes and glucosinolates as inducible physical andchemical resistance factors, respectively (Agrawal 1998, 1999). Likewise, in Nicotianaattenuata, the induced production of nicotine as a chemical resistance factorwas associated with metabolic costs, but provided a fitness benefit when plants wereunder attack by herbivores (Baldwin 1998; see also Steppuhn and Baldwin this volume).Although these findings should not be generalized and a defensive role shouldnot be assumed for all plant responses to wounding and herbivory, the prevalenceof inducible resistance traits in present day plant-herbivore systems implies thatsuch responses are likely the result of natural selection imposed by insect herbivoresduring evolution.
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