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oth as protective weapons and as potential signals. If <strong>this</strong> also holds f<strong>or</strong> chemical defenses, neighb<strong>or</strong> effects might have a<br />

_ndamental imp<strong>or</strong>tance in shaping plant defenses. This may well be so as, because acc<strong>or</strong>ding to Provenza et al. (1992),<br />

mmmalian herbiv<strong>or</strong>es learn to select food items through two interrelated systems. The affective system integrates the taste<br />

f food and its post-ingestive consequences, while the cognitive system integrates the od<strong>or</strong> and sight of food and its taste.<br />

tecause they have to sample plants in <strong>or</strong>der to adjust food intake to avoid intoxication (Provenza et al. 1992), the expected<br />

erbiv<strong>or</strong>y load of a plant is likely to depend both on its own defenses and on the defensive status of its neighb<strong>or</strong>s.<br />

SUMMARY<br />

In <strong>or</strong>der to be evolutionary stable, plant defenses should not benefit palatable neighb<strong>or</strong>s too much. If they do,<br />

_alatable plants can invade and eventually either out-compete unpalatable plants <strong>or</strong> establish polym<strong>or</strong>phic populations.<br />

Theref<strong>or</strong>e, we expect that plant defenses that can provide signals f<strong>or</strong> herbiv<strong>or</strong>es could be superi<strong>or</strong> to defenses that have no<br />

,alue as signals. A game the<strong>or</strong>etical analysis is presented in <strong>or</strong>der to expl<strong>or</strong>e the evolution of plant defenses when plant<br />

]tness depends on the defensive status of the neighb<strong>or</strong>.<br />

LITERATURE CITED<br />

\TSATT, RR.and O'DOWD, D.J. 1976. Plant defense guilds. Science 193: 24-29.<br />

_.UGNER, M., FAGERSTROM, T., and TUOMI, J. 1991. Competition, defense and games between plants. Behav. Ecol.<br />

Sociobiol. 29:231-234.<br />

3OMZE, I.M. 1983. Lotka-Volterra equation and replicat<strong>or</strong> dynamics: A two-dimensional classification. Biol. Cybernetics<br />

48:201-211.<br />

)ANELL, K., EDENIUS, L., and LUNDBERG, R 1991. Herbiv<strong>or</strong>y and tree stand composition: Moose patch use in winter.<br />

Ecology 72: 1350-1357.<br />

ESHEL, I. 1972. On the neighb<strong>or</strong> effect and the evolution of altruistic traits. The<strong>or</strong>. Pop. Biol. 3: 258-277.<br />

3UILFORD, T. and CUTHILL, I. 1991. The evolution of aposematism in marine gastropods. Evolution 45:449-451.<br />

HAY, M.E. 1986. Associational plant defenses and the maintenance of species diversity: Turning competit<strong>or</strong>s into accomplices.<br />

Am. Nat. 128: 617-641.<br />

HJALTEN, J., DANELL, K., and LUNDBERG, R I993. Herbiv<strong>or</strong>e avoidance by association: Vole and hare utilization of<br />

woody plants. Oikos 68: 125-131.<br />

HOFBAUER, J. and SIGMUND, K. 1988. The The<strong>or</strong>y of Evolution and Dynamical Systems. Cambridge Univ. Press,<br />

Cambridge.<br />

LAUNCHBAUGH, K.L. and PROVENZA, F.D. 1993. Can plants practice mimicry to avoid grazing by mammalian<br />

herbiv<strong>or</strong>es? Oikos 66: 501-504.<br />

MATTSON, W.J., HERMS, D.A., WITTER, J.A., and ALLEN, D.C. 1991. Woody plant grazing systems: N<strong>or</strong>th American<br />

outbreak foliv<strong>or</strong>es and their host plants, p. 53-84. In Baranchikov, Y., Mattson, W.J., Hain, ER, and Payne, T.L., eds.<br />

F<strong>or</strong>est insect guilds: patterns of interaction with host plants. Gen. Tech. Rep. NE-153. Radn<strong>or</strong> PA: U.S. Department<br />

of Agriculture, F<strong>or</strong>est Service. 400 p.<br />

McNAUGHTON, S.J. 1978. Serengeti ungulates: Feeding selectivity influences the effectiveness of plant defense guilds.<br />

Science 199: 806-807.<br />

NEE, S. 1989. Does Hamilton's rule describe the evolution of reciprocal altruism? J. The<strong>or</strong>. Biol. 141: 81-91. "<br />

37

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