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investigation. Sampling was done on 3 occasions at 2-week-intervals during needle expansion in spring: needles were taken from twigs of the upper crown region, where the sawflies prefer to oviposit. To record differences between the physiologically older needles at the bottom and the physiologically younger needles at the top of a given shoot, on the second and third occasion needles were separated according to their location on the expanding bud: top (t), middle (m), and bottom (b) needles. In contrast to stand A, the trees on stand B were undergoing an actual heavy sawfly attack during the sampling period in spring 1989. Additionally, large variation in the timing of bud burst of the trees was found: the early flushing individuals flushed 4 weeks before the late flushing ones. Sawfly attack, however, occurred only on the late flushing trees. The early flushing trees (n = 10) were harvested 3 times (1,2,3), the late flushing trees (n = 10) only twice (la, 2a). After the second sampling of the attacked, late flushing trees (2a), the larvae were fully grown and were no longer feeding. Although the sampling on early and late flushing trees differed by more than 4 weeks, needles from sampling date 1 corresponded phenologically with 1a and needles from sampling date 2 corresponded with 2a. A fertilization experiment was done on stand C with late flushing, attacked trees. The stand was fertilized with 130 kg nitrogen per ha/year, applied on 4 occasions: in summer 1989, in spring and summer 1990, and in spring 1991. Sampling was done during needle expansion in June, 1991, from 15 fertilized and 17 unfertilized (i.e., control) trees. Samples for chemical analyses were put into liquid nitrogen, then freeze-dried and milled. Nitrogen (total and soluble) was determined by micro-Kjeldahl technique with selenium as catalyst. Soluble carbohydrates (sugars, cyclitols) as well as the organic acids (quinic acid, shikimic acid) were determined by GLC, starch content was estimated enzymatically as glucose-equivalents. Needle phenolics were defined and determined as the potential of the needle extract (done with boiling 50% aqueous methanol) to precipitate bovine serum albumin (BSA). Absorption was measured at 590 nm. Fiber (cellulose, hemicellulose, lignin) was estimated after a method to determine the fiber content of cattle food: after hydrolyzation with strong acids and alkalines and heating up to 550 degrees, the remaining ash residue was weighed. All values are expressed on a dry weight basis. Larval experiments were performed at stand C with late flushing, fertilized, and unfertilized trees in spring 1991. Larvae from naturally attacked buds as well as larvae (lst to 3rd instar) placed on expanding buds and enclosed in cages were studied. After cocoon formation, larvae were collected, sexed, and weighed. Parasitized larvae were excluded from the analyses. RESULTS During initial feeding in mid-May (bud length 4 cm) the young sawfly larvae are confronted with both high total and soluble nitrogen (2% N tot. and 0.2% N sol.) and organic acid concentrations (12%). On the other hand, carbohydrates (sugars and cyclitols) and starch are at rather low levels (8% and 2%). Fiber was approximately 10% of the needle dry weight (dw). For all these traits, damaged and undamaged trees did not differ (Fig. 1, upper graph). The crucial difference, however, was the phenolics: concentrations were significantly lower in the attacked trees than in the unattacked trees (p
40 ........................................ i.......................... T.......................... i........................... [........................... '..........=_ ......":.......................... ! i LDa,fe lii;iiiiiiii i i cte 30 ............................. i ......................................................................................... ................... _' 20 ............................. "0 _ '!'!'!!!!!!!!!_!_!'! .............. ...... i 10 I ............ ..... i ]iiiii!i_iiiiii]ii_!_l iiEii::iiiiiiiiii!i!i I .... ,_i_,_,_,E _ ! !ii[::::iiii_ii!i_i!_ ........ , ii!iiiiii_i_iliiiiiii_i_l i!i!ii]i]iiii!]!iiii_;i! . ::ii_iiii_,i!i!i!i!i!i!i! 0 i iiiii;iiii_;i!iii::i::iiil I ' _........... ::: :'_'_ bud phenolics* total soluble soluble starch organic fiber expansion nitrogen nitrogen carbohydr, acids cm mg mg 0 ........................................................................................................................................................................................................................... 30 ............................................................................................................. ........................... _ ....................................... D affected _'20 .......................... _............. _.......................... _.......................... _........................... r.......................... i............................ i.... l°una ecte°] Io OIl!iil ........ ................ ............. bud expansion phenolics** total nitrogen soluble soluble starch organic acids fiber cm mg nitrogen mg carbohydrate Figure 1.--Spruce needle chemistry during initial (upper graph) and at the end (lower graph) of larval feeding on stand A (identical flushing time). Asterisks indicate significant differences between once affected (n = 15) and unaffected (n = 15)trees (levels of significance: *p < 0.05, ** p < 0.01). 250
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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
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HUNTER, M.D. and PRICE, P.W. 1992.
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A° ADDITIVE C. HYBRID SUSCEPTIBILI
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Galls, leaf miners, or leaf folds w
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Table 2.--Summary of the hypotheses
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the inheritance patterns of seconda
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FLOATE, K.D. and WHITHAM, T.G. 1993
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Elements of the Suitability/Defense
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Table l.--Comparing the mean number
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ACKNOWLEDGEMENTS The authors sincer
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SINGH, D.R 1986_ Breeding for resis
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The objectives of this presentation
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Table 1.--Average weevil attack on
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1977, Kline and Mitchell 1979, Wood
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Further research is underway to cla
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A SUGI CLONE HIGHLY PALATABLE TO HA
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RESULTS Seasonal Stability of Palat
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-
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OGAWA, A., NAGATA, Y., and SUKENO,
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MATERIALS AND METHODS Field Work In
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Laboratory Procedures The week afte
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E 120 Nogent-sur-Vernisson Remnings
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Interdependence Between Reaction Zo
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alance hypothesis (Lorio 1986) cann
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SOLHEIM, H., LANGSTROM, B., and HEL
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In a second experiment, three 30-ye
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Considering biochemical pathways, i
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- = 120 N /'_" =40 1/ _ j Days 0 _
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BIGGS, A.R. 1985. Suberized boundar
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DIFFERENTIAL SUSCEPTIBILITY OF WHIT
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Table 2.--Mortality of white fir pr
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Results from the geographic range p
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esulting seedlings were then rooted
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-0.2 0 5 82. _j -0,4 e a a a D4 E I
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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
Page 240 and 241: The importance of a compound as a r
Page 242 and 243: WERNER, R.A. 1995. Toxicity and rep
Page 244 and 245: 41 40 42 \ 4t 42 41 42 ",,,\ 42 40
Page 246 and 247: Table 1.--Summary data for the 26 p
Page 248 and 249: One weakness of using data from gen
Page 250 and 251: Lower Peninsula (populations 14-26)
Page 252 and 253: AUCLAIR, A.ND., WORREST, R.C., LACH
Page 254 and 255: KELLOMAKI, S., HANNINEN, H., and KO
Page 256 and 257: WARGO, RM. and HAACK, R.A. 1991. Un
Page 260 and 261: Figure 2.--Changes in protein preci
Page 262 and 263: q) J 400 T J ! i i i i o control tr
Page 264 and 265: attacked trees than in control tree
Page 266 and 267: ACIDIC DEPOSITION, DROUGHT, AND INS
Page 268 and 269: Table 1.--Impact of acidic fog upon
Page 270 and 271: MENGEL, K., BREININGER, T., and LUT
Page 272 and 273: THE RESISTANCE OF SCOTCH PINE TO DE
Page 274 and 275: EXPERIMENTAL METHODS Experiments we
Page 276 and 277: C w Q. 1,200 1,000 o 800 C_ --- 600
Page 278 and 279: 0 .................................
Page 280 and 281: FONTAtNE, R.G. 1985. Forty years of
Page 282 and 283: Host Defenses An important componen
Page 284 and 285: Functional Heterogeneity of Forest
Page 286 and 287: The Connectivity Index (CI). The CI
Page 288 and 289: FUNCTIONAL HETEROGENEITYANALYSIS :
Page 290 and 291: The Modified Hazard Map. The next l
Page 292 and 293: systems facilitate mapping of fores
Page 294 and 295: KOLASA, J. and ROLLO, C.D. 1991. In
Page 296: ) i,i_i)i_)i_i_ U.S. Departme_]t of
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