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Induced Plant Responses to Herbivory - Terrestrial Systems Ecology

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338 KARBAN & MYERS<br />

Annu. Rev. Ecol. Syst. 1989.20:331-348. Downloaded from www.annualreviews.org<br />

by ETH- Eidgenossische Technische Hochschule Zurich - BIBLIOTHEK on 03/29/11. For personal use only.<br />

Many ecologists became interested initially in induced responses because<br />

they provided a potential mechanism <strong>to</strong> explain multiyear population cycles<br />

of forest insects. The hypothesis presented by Haukioja & Hakala (38) and<br />

Benz (8)-that plant quality decreases after defoliation and then increases<br />

gradually after a lag of several years-provides a delayed density-dependent<br />

mechanism that could potentially drive population cycles of herbivores (11,<br />

36, 87).<br />

To explain regional synchrony of population fluctuations of forest Lepidoptera,<br />

we must test whether host trees respond in a consistent manner <strong>to</strong> insect<br />

attack. This basic premise does not seem <strong>to</strong> be supported: <strong>Induced</strong> responses<br />

of trees have been found <strong>to</strong> vary among species, among populations, among<br />

years, and across environmental gradients (81a). On the other hand, changes<br />

in the fecundity and survival of fluctuating populations of forest Lepidoptera<br />

often show consistent patterns through the cycle, even when caterpillars feed<br />

on different species of host plant, in different areas, and following different<br />

his<strong>to</strong>ries of attack (77, 78).<br />

Although the variation in response of trees <strong>to</strong> herbivore damage seems <strong>to</strong><br />

make inducible changes in food quality an unlikely explanation for the cyclic<br />

population dynamics of forest Lepidoptera, we list in Table 2 further predictions<br />

of the hypothesis that can be tested. Observations on cyclic populations<br />

of tent caterpillars and other forest Lepidoptera do not support<br />

these predictions (77, 78). The importance of inducible changes in food<br />

plant quality <strong>to</strong> population dynamics of nonoutbreak species has not been<br />

studied.<br />

Table 2<br />

Testable predictions arising from the hypothesis that population cycles of forest<br />

Lepidoptera are driven by deterioration in food plant quality following feeding damage from<br />

increasing numbers of herbivores. Species and populations of host trees must respond in a<br />

consistent manner <strong>to</strong> herbivore damage for the fluctuations of different populations of insects <strong>to</strong><br />

remain in synchrony within a region.<br />

1. Fecundity and survival of herbivores will be related <strong>to</strong> the his<strong>to</strong>ry of attack on trees.<br />

2. If the response of trees is density dependent, fecundity and survival of herbivores will decline<br />

with increasing density (level of attack) and deterioration in food quality.<br />

3. Decreasing fecundity and survival of herbivores following damage <strong>to</strong> host plants will be<br />

translated in<strong>to</strong> a decline in the population density.<br />

4. Cropping of herbivore density <strong>to</strong> reduce damage will prolong the outbreak phase of the<br />

population.<br />

5. Introduction of herbivores <strong>to</strong> suitable foodplants in sites with no previous herbivore damage<br />

will lead <strong>to</strong> an outbreak out of synchrony with natural populations.

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