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Considering biochemical pathways, it is interesting to note that the appearance of stilbenes and Pc in Scots pine corresponded to decreases in compounds of the hydroxycinnamic acid group, since this latter group is a precursor of both stilbenes and flavonoids (Ribereau-Gayon 1968, Gotham 1989). Between Clone Comparisons Clonal variability was first analyzed by multivariate analysis, in order to take into account all characterized compounds together. A canonical discriminant analysis was performed by taking reaction zones as Inain individuals and unwounded phloem as complementary individuals. This procedure was necessary as the purpose was mainly to compare clone :reaction zones with each other, and only to place unwounded phloem relatively to them. Axes 1 2 and 3 explained 57, 23, and 15% of the variation, respectively. In the first two axes (Fig. 3), froln un- ' wounded phloem to the reaction zone, phenolic composition seemed to evolve towards the same point of the plane, whatever the clone. That was confirmed in all significant axes (three), by calculating mean distances between points and their barycenter. Spread of the points relating to reaction zones was less than spread of the points relating to unwounded phloeln, in each clone and for all clones together (data not shown). Thus, each clone would react in a different way according to its constitutive composition, but always so that the composition of the reaction zone tended to be the same. The meaning of this phenolic modifications is not clear. It may be linked with protection against aggressors or with tissue restoration. Nevertheless, it is worthwhile to note that clone ...... c , whose reaction was the least efficient (reaction zone significantly longer than that of' all other clones), was the only one initially separated from the others by axis 2 (Fig. 3). If phenols effectively play a role in arresting beetles and their fungi, it could thus be possible to predict reaction efficiency of a clone on the basis of the whole phenolic composition of its constitutive phloem. 8 _'.%0 eql -8 . d ""N& 2 3, 4 4 / 5 b'b_za 6 "_so_ s I I I II -10 0 10 Axis 1 (57%) Figure 3._Canonical discriminant analysis of phloem phenolic composition of different Scots pine clones, after inoculation with Ophiostoma brunneo-ciliatum. Reaction zones were considered as principal individuals and unwounded phloem as complementary individuals, a, b, c, d, e, f = unwounded phloem of the different clones (* represents the barycenters); 1, 2, 3, 4, 5, 6 = Corresponding reaction zones (o represent the barycenters). Arrows indicate the directions of variation from unwounded phloems to reactions zones. Clones a, b, c, d, e and f were respectively represented by 3, 2, 1, 2, 3 and 3 trees. t81
Clonal variability was also analyzed compound by compound, for the peaks characterized at least at the family level. To appreciate if certain compounds could be used as markers of efficiency of tree response, phenol concentration in each clone and for each compound, was compared to reaction zone length. The best correlation (r = 0.78) was for taxifolin glycoside in unwounded phloem (data not shown). The lowest concentrations corresponded to the least extended reaction zones, that is to the most effective response, and inversely. Concentration of taxifolin glycoside could thus be a marker of tree's response efficiency. Presence of taxifolin glycoside in the phloem prior to aggression could be favorable to the fungus, or could directly impede the development of the tree response. In the two cases, the result would be an increase in the lag time necessary to stop the fungus, thus causing a long reaction zone. Kinetics of the Phenolic Response A first approach to the kinetics was performed with multivariate analysis, the same way as in the previous experiment (Fig. 4). Axes 1 and 2 explained 88 and 9 percent of the variation, respectively. The composition of all unwounded samples was close to that of reaction zones at day 3, which suggest that phenolic composition of unwounded phloem did not vary 3 1 4 u ,,_,,, ..t 4 2 3 4 333 5 S 2434 5 At ed s -3 d [ -3 ' -i " i ' ; '.... i ' Axis 1 (88%) Figure 4.--Canonical discriminant analysis of the phenolic composition of Scots pine phloem collected at different dates after inoculations with Ophiostoma brunneo-ciliatum, in three different trees. Reaction zones were considered as principal individuals and unwounded phloem as complementary individuals. 1, 2, 3, 4, 5 = Reaction zones collected respectively 3, 7, 14, 30 and 60 days after inoculations. A, B, C, D, E = corresponding barycenters, d, e = unwounded phloem collected at days 30 and 60. Arrows indicate the directions of variation during reaction development. during the whole experiment. Although between tree variability existed, the phenolic composition of the reaction zones at day 60 was clearly separated from that of all other samples along axis 1. Reaction zones at days 3 were separated from other samples along axis 2. Kinetics of phenolic response thus proceeded in two main phases. Concomitantly, growth of the reaction zone also proceeded in two main phases and was approximately done at day 30, indicating that the fungus had been stopped before this date (Fig. 5). One can thus hypothesize that the first phase, before day 30, is related to resistance and to interactions between tree and fungus, while the second more probably corresponds to wound healing processes. Considenng that, it is interesting to note that wound periderm began to be visible at day 30. Analysis, compound by compound, revealed three main kinds of kinetics. Ps, Psme and Pc increased in two phases, first early until day 7, then late after day 30 (Fig. 6a and 6b). Some other compounds, such as acetophenone glycoside, regularly decreased or remained constant (Fig. 6c), while others, such as taxifolin, increased only late, after day 30, SOmetimes after a decrease at the very beginning of development of the reaction (Fig. 6d). 182 5 , i: i'i:i_i_'i!ill
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DYNAMICS OF FOREST HERBIVORY: QUEST
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TABLE OF CONTENTS Seeking The Rules
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32. Companion planting of the nitro
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GRIMALSKY, ¥.I., Research Institut
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POUTTU, ANTTI, Finnish Forest Resea
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persist in mature leaves and affect
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THE SINK-SOURCE HYPOTHESIS: TYPE AN
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late season defoliation decreases t
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BLANCHE, C.A., LORIO, EL., Jr., SOM
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NIEMELA, R, TUOMI, J,, and LOJANDER
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induced reactions have been studied
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summer, but the other sawfly specie
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LARSSON, S., BJ(_RKMAN, C., and GRE
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STAND AND LANDSCAPE DIVERSITY AS A
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not related to resistance to the se
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In contrast to the situation in eas
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ROOT, R.B. 1973. Organization of a
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We tlhus neglect two potential sour
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c__ c__ 1T1 Ill a) a) 0 0 0 e 1 o e
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c_ >l.s S S S m N < DN DN < D c DN
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:hen k > 0 suggest that the cue or
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oth as protective weapons and as po
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DEFENSE THEORIES AND BIRCH RESISTAN
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P 45 A L 40 A T 35 A B 30 I L 25 I
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However, hares showed strong prefer
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DUGLE, J.A. 1966. A taxonomic study
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accommodate findings related to mic
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Table l.--Mean ±S.E. area eaten by
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Sulfhydryl-Disulfide Mechanisms In
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Iraa prior experimental analysis of
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FRAZIER, J.L. and HEITZ, J.R. 1975.
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REICHARD, R 1993. From RNA to DNA,
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; trees, and between and within ind
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Encapsulated objects are data struc
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_re3.--The Lignum model can be cont
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SEVERE DEFOLIATION Tree Fine Root +
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RUMBAUGH, J., BLAHA, M., PREMERLANI
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oo0 a b ol ¢' _achiman_i ()® 1,eA
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1000 Conspicuous Defo i at ion 100
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2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
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Qualitative Changes in [,eaves and
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t00 2 Yr 1 Yr o_ 80 Treatment Treat
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Table 2.--Body size of Q. purzctate
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06 b AddedBSA < 04 _ 2rag -'=' i:3
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Severe Defol iat ion Site-A 1993 _
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KAMATA, N. and IGARASHI, M. 1995b.
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enclosure %r 20 post-diapausing sec
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2,500 F 000 F NF NF b _7 a _;_ a _7
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8O D 09 i 4,'.) F AS4L a Z___7 2o !
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2,500 _ st2-st4 El st5 0 st6 [_ pup
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96 SLANSKY, E, Jr. 1990. Insect nut
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later, we selected one more floweri
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Table 2.---Mean spruce budworm perf
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Table 3..--Mean spruce budworm perf
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FOLIVORE FEEDING ON MALE CONIFER FL
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Table 3.--Lymantria monacha prefere
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500- 400- i 300 v ¢ m a 200- I00 L
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From the point of view of trees, th
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THE BLACKMARGINED APHID AS A KEYSTO
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Wood et al. (1987) report that M. c
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MIZELL, R.E 1991. Pesticides and be
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METHODS Study Site The study took p
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the samples. Fertilization had subs
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Table 2.--Percent nutrient content
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Some tree responses to fertilizatio
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PHYTOCHEMICAL PROTECTION AND NATURA
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Indirect effects of abiotic factors
Page 140 and 141: HUNTER, M.D. and PRICE, P.W. 1992.
Page 142 and 143: A° ADDITIVE C. HYBRID SUSCEPTIBILI
Page 144 and 145: Galls, leaf miners, or leaf folds w
Page 146 and 147: Table 2.--Summary of the hypotheses
Page 148 and 149: the inheritance patterns of seconda
Page 150 and 151: FLOATE, K.D. and WHITHAM, T.G. 1993
Page 152 and 153: Elements of the Suitability/Defense
Page 154 and 155: Table l.--Comparing the mean number
Page 156 and 157: ACKNOWLEDGEMENTS The authors sincer
Page 158 and 159: SINGH, D.R 1986_ Breeding for resis
Page 160 and 161: The objectives of this presentation
Page 162 and 163: Table 1.--Average weevil attack on
Page 164 and 165: 1977, Kline and Mitchell 1979, Wood
Page 166 and 167: Further research is underway to cla
Page 168 and 169: A SUGI CLONE HIGHLY PALATABLE TO HA
Page 170 and 171: RESULTS Seasonal Stability of Palat
Page 172 and 173: -
Page 174 and 175: OGAWA, A., NAGATA, Y., and SUKENO,
Page 176 and 177: MATERIALS AND METHODS Field Work In
Page 178 and 179: Laboratory Procedures The week afte
Page 180 and 181: E 120 Nogent-sur-Vernisson Remnings
Page 182 and 183: Interdependence Between Reaction Zo
Page 184 and 185: alance hypothesis (Lorio 1986) cann
Page 186 and 187: SOLHEIM, H., LANGSTROM, B., and HEL
Page 188 and 189: In a second experiment, three 30-ye
Page 192 and 193: - = 120 N /'_" =40 1/ _ j Days 0 _
Page 194 and 195: BIGGS, A.R. 1985. Suberized boundar
Page 196 and 197: DIFFERENTIAL SUSCEPTIBILITY OF WHIT
Page 198 and 199: Table 2.--Mortality of white fir pr
Page 200 and 201: Results from the geographic range p
Page 202 and 203: esulting seedlings were then rooted
Page 204 and 205: -0.2 0 5 82. _j -0,4 e a a a D4 E I
Page 206 and 207: 1960, Kalkstein 1976). Increased su
Page 208 and 209: RYAN, B.F., JOtNER, B.L., and RYAN,
Page 210 and 211: It is seldom feasible to control wa
Page 212 and 213: 15 15 u. 10 |O O N tal 5 5 g ao o 4
Page 214 and 215: Only recently have we conducted stu
Page 216 and 217: esistance to beetle attack. Another
Page 218 and 219: LORIO, EL., Jr'. and HODGES, J.D. 1
Page 220 and 221: EFFECTS OF ROOT INHABITING INSECT-F
Page 222 and 223: OCCURRENCE OF ROOT AND STEM SUBCORT
Page 224 and 225: ]Predisposition of Root-infected Tr
Page 226 and 227: chemistry associated with root colo
Page 228 and 229: Implications to Natural Resource Ma
Page 230 and 231: KLEPZIG, K.D., RAFFA, K.F., and SMA
Page 232 and 233: RAFFA, K.F., PHILLIPS, T., and SALO
Page 234 and 235: METHODS The study was installed in
Page 236 and 237: FURNISS, M.M., LIVINGSTON, R.L., an
Page 238 and 239: METHODS In the spring and summer of
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The importance of a compound as a r
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WERNER, R.A. 1995. Toxicity and rep
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41 40 42 \ 4t 42 41 42 ",,,\ 42 40
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Table 1.--Summary data for the 26 p
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One weakness of using data from gen
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Lower Peninsula (populations 14-26)
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AUCLAIR, A.ND., WORREST, R.C., LACH
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KELLOMAKI, S., HANNINEN, H., and KO
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WARGO, RM. and HAACK, R.A. 1991. Un
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investigation. Sampling was done on
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Figure 2.--Changes in protein preci
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q) J 400 T J ! i i i i o control tr
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attacked trees than in control tree
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ACIDIC DEPOSITION, DROUGHT, AND INS
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Table 1.--Impact of acidic fog upon
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MENGEL, K., BREININGER, T., and LUT
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THE RESISTANCE OF SCOTCH PINE TO DE
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EXPERIMENTAL METHODS Experiments we
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C w Q. 1,200 1,000 o 800 C_ --- 600
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0 .................................
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FONTAtNE, R.G. 1985. Forty years of
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Host Defenses An important componen
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Functional Heterogeneity of Forest
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The Connectivity Index (CI). The CI
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FUNCTIONAL HETEROGENEITYANALYSIS :
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The Modified Hazard Map. The next l
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systems facilitate mapping of fores
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KOLASA, J. and ROLLO, C.D. 1991. In
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) i,i_i)i_)i_i_ U.S. Departme_]t of
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