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CAN PHLOEM PHENOLS BE USED AS MARKERS OF SCOTS PINE<br />

RESISTANCE TO BARK BEETLES?<br />

F. LIEUTIER _,F. BRIGNOLAS l, V. PICRON t, A. YART l, and C. BASTIEN 2<br />

t<strong>Station</strong> de Zoologie F<strong>or</strong>esti_re, Institut National de la Recherche Agronomique<br />

Ardon, 45 160 Olivet, France<br />

2<strong>Station</strong> d'Am61i<strong>or</strong>ation des Arbres F<strong>or</strong>estiers, Institut National de la Recherche Agronomique<br />

Ardon, 45 160 Olivet, France<br />

INTRODUCTION<br />

Bark beetle damage is considerable in all coniferous f<strong>or</strong>ests in the temperate zone. In most cases, it results from<br />

beetle attacks surpassing the critical density (numbers/m 2)below which trees can successfully reject them. Above <strong>this</strong><br />

threshold, natural mechanisms of tree resistance are overcome by the beetles and fungi which become successfully established.<br />

The level of <strong>this</strong> critical threshold of attack density is a fundamental expression of the strength <strong>or</strong> capacity of tree<br />

resistance (Berryman 1978, 1982; Christiansen et al. 1987).<br />

The basic phenomenon involved in the resistance is a phloem reaction induced by the invaders and localized around<br />

them. It consists of a rapid synthesis of chemicals, including secondary metabolites such as terpenes and phenols, which<br />

impregnate tree tissues (Reid et al. 1967, Berryman 1972, Shrimpton 1973, Christiansen and H<strong>or</strong>ntvedt 1983, Christiansen et<br />

al. 1987, Lieutier et al. 1988, Lfmgstr6m et al. 1992, Lieutier 1993). The critical threshold of attack density depends on the<br />

quantity of energy the tree is able to rapidly mobilize f<strong>or</strong> various syntheses in the reaction zones, and <strong>this</strong> quantity is necessarily<br />

limited (Christiansen et al. 1987). Thus, the threshold can be high if the lag time f<strong>or</strong> stopping an aggress<strong>or</strong> at each<br />

point of attack is very sh<strong>or</strong>t and hence the energy mobilized f<strong>or</strong> that purpose in each reaction zone is low. Under these<br />

conditions of arrested attacks, a sh<strong>or</strong>t reaction zone has been assumed to be m<strong>or</strong>e effective than an extended one (Lieutier et<br />

al. 1993).<br />

Markers of conifer resistance to bark beetles could be very useful; e.g., f<strong>or</strong>ecasting susceptibility to beetle attacks and<br />

beetle outbreaks, and aiding in selection of highly resistant trees. Because the concentrations of secondary metabolites vary<br />

considerably during tree responses to aggression, they merit a special interest. Preliminary studies on phloem phenolics have<br />

suggested dramatic changes in Scots pine, Pinus sylvestris L., in response to aggression (Lieutier et al. 1991a). This paper<br />

presents the results from two complementary approaches recently developed in France concerning the possible role of<br />

phenols in arresting bark beetle invasion and their use as biochemical markers of resistance. Detailed methods and other<br />

findings are presented elsewhere (Lieutier et al. 1995).<br />

METHODS<br />

Experimental Devices<br />

In our first experiment, six clones located in the INRA's nursery in Orlfians (France), each represented by one to three<br />

10-year-old trees, were used to test if a relationship existed between reaction zone length and phenolic composition. Trees<br />

were each inoculated with a malt agar sp<strong>or</strong>ulated culture of Ophiostoma brunneo-ciliatum Math.-K., a fungus previously<br />

isolated from lps sexdentatus Boern., and thought to play a role in the establishment of <strong>this</strong> beetle in Scots pine (Lieutier et<br />

al. 1988). Each tree received three inoculations. Three weeks later, the reaction zone around each inoculation point was<br />

measured and sampled. Three samples of unwounded phloem were also taken from each tree.<br />

Mattson, W.J., Niemel_i, P., and Rousi, M., eds. 1996. Dynamics of f<strong>or</strong>est herbiv<strong>or</strong>y: quest f<strong>or</strong> pattern and principle.<br />

<strong>USDA</strong> F<strong>or</strong>. Serv. Gen. Tech. Rep. NC-183, N.C. F<strong>or</strong>. Exp. Sta., St. Paul, MN 55108.<br />

178

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