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attacked trees than in control trees. Successful larval feeding seems to be only possible on trees with low needle phenolics. High phenolic concentrations during early larval feeding suggest an effective defense mechanism of the host tree and seems to be genetically determined. A significant negative correlation between needle nitrogen and phenolics in individual trees as well as the positive effect of nitrogen and the negative effect of phenolics on larval weight indicate a causal relationship between nitrogen and phenolics and larval growth. We also demonstrated the effects of excess atmospheric nitrogen input via N fertilization upon different chemical properties in spruce needles and the quality of these expanding needles as food for the larvae. Nitrogen fertilization resulted in decreased phenolics, starch, organic acids, and fiber, increased nitrogen and carbohydrates, and in a better food value for the sawfly larvae. Larval weight was negatively correlated with needle phenolic concentrations. Our findings indicate that the quality of spruce needles as food for the P abietina larvae are influenced in two ways: by increasing the concentrations of positive factors (e.g., nitrogen, carbohydrates) and decreasing the concentrations of negative factors (e.g., phenol ics, fiber, organic acids). ACKNOWLEDGEMENTS This study was funded by the Austrian Fonds of Scientific Research, P 7093-CHE: Excess Nitrogen in Forest Plant-Herbivore Associations. The authors wish to thank Ing. G. Mottlik, A. Stradner, and Mag. A. Straka for technical assistance with the chemical analyses. LITERATURE CITED BALSBERG-PAHLSSON, A.M. 1992. Influence of nitrogen fertilization on minerals, carbohydrates, amino acids and phenolic compounds in beech (Fagus sylvatica L.) leaves. Tree Physiol. 10: 93-100. BERGER, R. 1992. Massenauftreten der Kleinen Fichtenblattwespe Pristiphora abietina (Christ) im Hausruck. Teil I: Okologische Rahmenbedingungen in einem "untypischen" Befallsgebiet. Anz. Sch_idlingskde., Pflanzenschutz, Umweltschutz 65: 105-114. BERGER, R. and KATZENSTEINER, K. 1994. Massenauftreten der Kleinen Fichtenblattwespe Pristiphora abietina (Christ) im Hausruck. Teil II: Immissions6kologische Einfltisse. J. appl. Ent. (in press). BRYANT, J.E, CHAPIN, F.S., KLEIN, D.R. 1983. Carbon/nutrient balance of boreal plants in relation to vertebrate herbivory. Oikos 40: 357-368. BRYANT, J.E, CLAUSEN, T.E, REICHARDT, EB., McCARTHY, M.C., and WERNER, R.A. 1987. Effect of nitrogen fertilization upon the secondary chemistry and nutritional value of quaking aspen (Populus tremuloides Michx.) leaves for the large aspen tortrix (Choristoneura conflictana (Walker)). Oecologia 73:513-517. COLEY, ED. 1983. Herbivory and defensive characteristics of tree species in a lowland tropical forest. Ecol. Monogr. 53: 209-233. ERICSSON, A. 1979. Effects of fertilization and irrigation on the seasonal changes of carbohydrate reserves in different age-classes of needle on 20-year-old Scots pine trees (Pinus sylvestris). Physiol. Plant. 45: 270-280. FEENY, EE 1970. Seasonal changes in oak leaf tannins and nutrients as a cause of spring feeding by winter moth caterpillars. Ecology 51:565-581. FEENY, EE 1976. Plant apparency and chemical defense. Recent Adv. Phytochem. 19: 1-40. 255
GLYPHIS, J.R and PUTTICK, G.M. 1989. Phenolics, nutrition and insect herbivory in some garrigue and maquis plant species. Oecologia 78: 259-263. HATCHER, RE. 1990. Seasonal and age-related variation in the needle quality of five conifer species. Oecologia 85: 200- 212. HAUKIOJA, E., NIEMELA, R, and SIREN, S. 1985. Foliage phenols and nitrogen in relation to growth, insect damage, and ability to recover after defoliation, in the mountain birch Betula pubescens ssp tortuosa. Oecologia 65:214-222. HOLOPAINEN, J.K., KAINULAINEN, E., OKSANEN, J., WULFF, A., and KARENLAMPI, L. 1991. Effect of exposure to fluoride, nitrogen compounds and SO 2on the numbers of spruce shoot aphids on Norway spruce seedlings. Oecologia 86: 51-56. HOLZER, K. and SCHULTZE, U. 1988. Die Bedeutung yon Herkunft und Austrieb bei einem Fichtenblattwespenbefall. Cbl. ges. Forstwesen 105:207-216. JENSEN, T.S. 1988. Variability of Norway spruce (Picea abies L.) needles; performance of spruce sawflies (Gilpinia hercyniae Htg.). Oecologia 77:313-320. LUNDERST,ADT, J. 1980. Zur 6kophysiologischen Bedeutung yon Phenolen und Proteinen in Nadeln der Fichte (Picea abies Karst.). Z. Pflanzenern_ihr. Bodenkd. 143: 412-421. MATTSON, W.J. 1980. Herbivory in relation to plant nitrogen content. Ann. Rev. Ecol. Syst. 11: 119-161. MAUFFETTE, Y. and OECHEL, W.C. 1989. Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica. Oecologia 79: 439-445. NASHOLM, T. and ERICSSON, A. 1990. Seasonal changes in amino acids, protein and total nitrogen in needles of fertilized Scots pine trees. Tree Physiol. 6: 267-281. PUTTICK, G.M. 1986. Utilization of evergreen and deciduous oaks by the Californian oak moth, Phryganidia californica. Oecologia 68: 589-594. SCHAFELLNER, C., BERGER, R., MATTANOVICH, J., and FOHRER, E. 1993. Pristiphora abietina (Hym., Tenthredinidae) - ein Bioindikator ftir Luftverschmutzung? Besonderheiten unreifer Fichtennadeln als Larvenfutter. Forstw. Cb. 112: 116-128. SCHAFELLNER, C., BERGER, R., MATTANOVICH, J., and Ff0HRER, E. 1994. Food quality of spruce needles and the performance of the Little Spruce Sawfly, Pristiphora abietina Christ (Hym., Tenthredinidae). The protein precipitating ability of the young needles. Acta horticulturae (in press). SCHOPE R. 1986. The effect of secondary needle compounds on the development of phytophagous insects. For. Ecol. Man. 15: 55-64. SCRIBER, J.M. and SLANSKY, E 1981. The nutritional ecology of immature insects. Ann. Rev. Entomol. 26: 183-211. SIERPINSKI, Z. 1985. Luftverunreinigungen und Forstsch_idlinge. J. appl. Ent. 99: 1-6. TUOMI, J., NIEMEL,_, R, and SIREN, S. 1990. The Panglossian paradigm and delayed inducible accumulation of foliar phenolics in mountain birch. Oikos 59: 399-410. WARING, R.H., McDONALD, A.J.S., LARSSON, S., ERICSSON, T., WIREN, A., ARWIDSSON, E., ERICSSON, A., and LOHAMMAR, T. 1985. Differences in chemical composition of plants grown at constant relative growth rates with stable mineral nutrition. Oecologia 65: 157-160. 256
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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
Page 15 and 16:
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
Page 38 and 39:
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
Page 56 and 57:
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
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RYAN, B.F., JOtNER, B.L., and RYAN,
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It is seldom feasible to control wa
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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 258 and 259: investigation. Sampling was done on
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 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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