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GENETIC AND PHENOTYPIC VARIATION IN THE INDUCED REACTION OF<br />
SCOTS PINE TO LEPTOGRAPHIUM WINGFIELDII:<br />
REACTION ZONE LENGTH AND FUNGAL GROWTH<br />
F. LIEUTIER 1,B. LANGSTROM z, H. SOLHEIM 3<br />
C. HELLQVIST 2, and A. YART l<br />
_<strong>Station</strong> de Zoologie F<strong>or</strong>esti_re, Institut National de la Recherche Agronomique, Ardon, 45 160 Olivet, France<br />
2Division of F<strong>or</strong>est Entomology, Swedish University of Agricultural Sciences, S-77073 Garpenberg, Sweden<br />
3Department of F<strong>or</strong>est Ecology, N<strong>or</strong>wegian F<strong>or</strong>est <strong>Research</strong> Institute, As-NLH, N 1432, N<strong>or</strong>way.<br />
INTRODUCTION<br />
The induced defensive reaction of phloem generally plays a decisive role in the resistance of conifers to attack by<br />
bark beetles and their associated fungi. It is thought to be induced by fungi (Reid et al. 1967, Berryman 1972, Raffa and<br />
Berryman 1983, Christiansen and H<strong>or</strong>ntvedt 1983, Matson and Hain 1985, Christiansen et al. 1987, Lfingstr6m et al. 1992),<br />
<strong>or</strong> by the b<strong>or</strong>ing activity of the beetle (Lieutier et al. 1988, Lieutier 1993). In most cases, by stimulating that energy demanding<br />
response, the fungi can lower the threshold of attack density above which the tree is overwhelmed and beetle attacks are<br />
successful. In all situations, the individual reaction is the basic mechanism of tree resistance.<br />
The between-tree variability in the threshold of attack density (Waring and Pitman 1980, Mulock and Christiansen<br />
1986), as well as the between-tree variability in the induced reaction itself (Shrimpton and Reid 1973, Peterman 1977,<br />
Lieutier and Ferrell 1989, Lieutier et al. 1993), has been linked to tree vig<strong>or</strong>. However, the genetic contribution to <strong>this</strong><br />
variability has never been studied.<br />
In Scots pine, Pinus sylvestris L., an induced reaction is essential to resist attacks by lps sexdentatus Boern. and<br />
Tomicus piniperda L., and their associated fungi (Lieutier et al. 1988, 1989a; Solheim and L_ngstr6m, 1991; L_ngstr_m et al.<br />
1992). The blue-staln fungus, Leptographium wing_i'eldii M<strong>or</strong>elet, associated with T. piniperda, does not seem to play a role<br />
in the development of the induced reaction and in the success of attacks (Lieutier et al. 1988, 1989b). However, artificial<br />
inoculation of its sp<strong>or</strong>ulated cultures into Scots pine always incites strong defensive reactions (Lieutier et al. 1989a, Solheim<br />
and L_ngstr6m 1991) and the tree's response depends on the number of sp<strong>or</strong>es present in the inoculum (Lieutier et al. 1989b).<br />
Additionally, it may play a role in tree m<strong>or</strong>tality, since weakened Scots pine can be killed by a high density of artificial<br />
inoculations (Solheim et al. 1993).<br />
The present study is part of a cooperative project with the following objectives: (1) to study the variation in the<br />
induced reactions (reaction zone f<strong>or</strong>mation, sapwood occlusion) between Scots pine provenances in two different climatic<br />
regions, (2) to study the perf<strong>or</strong>mance of a French and a Swedish strain of L. wingfieldii in these provenances under different<br />
climatic conditions, and (3) to study the defensive chemistry involved in the induced reactions, particularly the roles of resin<br />
acids and phenolics, and to search f<strong>or</strong> chemical markers f<strong>or</strong> resistance among these groups of chemicals. This study addresses<br />
the first two issues.<br />
Mattson, W.J., Niemelfi, 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-I83, N.C. F<strong>or</strong>. Exp. Sta., St. Paul, MN 55108.<br />
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