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Annu. Rev. Entomol. 2006.51:45-66. Downloaded from arjournals.annualreviews.org<br />

by University of British Columbia Library on 12/09/05. For personal use only.<br />

56 ISMAN<br />

face the use of antifeedants <strong>in</strong> agriculture (47). The first is <strong>in</strong>terspecific variation<br />

<strong>in</strong> response—even closely related species can differ dramatically <strong>in</strong> behavioral<br />

responses to a substance—limit<strong>in</strong>g the range of pests affected by a particular antifeedant<br />

(43). Some substances that deter feed<strong>in</strong>g by one pest can even serve as<br />

attractants or stimulants for other pests. The second is the behavioral plasticity <strong>in</strong><br />

<strong>in</strong>sects—pests can rapidly habituate to feed<strong>in</strong>g <strong>deterrents</strong>, render<strong>in</strong>g them <strong>in</strong>effective<br />

<strong>in</strong> a matter of hours. This has been recently demonstrated not only for pure<br />

substances like azadiracht<strong>in</strong> (13), but also for complex mixtures (plant extracts)<br />

(1). Whereas a highly mobile (fly<strong>in</strong>g) <strong>in</strong>sect may leave a plant upon first encounter<strong>in</strong>g<br />

an antifeedant, a less mobile one (larva) may rema<strong>in</strong> on the plant long enough<br />

for the deterrent response to wane. Such behavioral changes are important <strong>in</strong> light<br />

of the observation that some plant substances are <strong>in</strong>itially feed<strong>in</strong>g <strong>deterrents</strong> but<br />

lack toxicity if <strong>in</strong>gested. Azadiracht<strong>in</strong> is clearly an exception to this rule, as <strong>in</strong>gestion<br />

leads to deleterious physiological consequences, but many other compounds<br />

or extracts with demonstrated antifeedant effects lack toxicity when adm<strong>in</strong>istered<br />

topically or via <strong>in</strong>jection (8, 9).<br />

Repellents<br />

For many chemists, an effective alternative to DEET (N,N-diethyl-m-toluamide)<br />

for personal protection aga<strong>in</strong>st mosquitoes <strong>and</strong> bit<strong>in</strong>g flies is the holy grail. In<br />

spite of five decades of research, no chemical has been found that provides the<br />

degree of protection aga<strong>in</strong>st bit<strong>in</strong>g mosquitoes or persistence on human sk<strong>in</strong> afforded<br />

by DEET (74). Concerns with the safety of DEET, especially to children,<br />

have resulted <strong>in</strong> the <strong>in</strong>troduction of several plant oils as natural alternatives. Some<br />

personal <strong>repellents</strong> <strong>in</strong> the U.S. marketplace conta<strong>in</strong> oils of citronella, eucalyptus,<br />

or cedarwood as active <strong>in</strong>gredients; 2-phenethylpropionate, a constituent of<br />

peanut oil, <strong>and</strong> p-menthane-3,8-diol (obta<strong>in</strong>ed from a particular species of m<strong>in</strong>t)<br />

(Figure 2) are also used <strong>in</strong> consumer products. All of these materials can provide<br />

some protection, but the duration of their effect can be limited (often

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