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However, hares showed strong preference for it. The general trend in our experiments has been that species and genotypes resistant to hares showed better than average growth, although the relationship was not statistically significant (Figs. 3 and 4). These results support our earlier findings which indicate either a positive relationship between first year growth of birch seedlings and their hare resistance, or no relationship at all (Rousi et al. 1989, 1990, 1991, 1996b). European white birch genotypes have not shown any relationship between their growth and resistance to hares, Microtus voles, Curculionidae weevils, or Melampsoridium leaf rust (Poteri and Rousi 1996, Rousi et al. 1996b). These experiments did not support any ecological trade-off either: resistance to any one pest was not related to resistance to others. However, vole and hare resistance in some experiments have shown a positive relationship (Chiba and Nagata 1969), and have also tended to do so in our own experiments (Rousi et al. t989, 1996b). Interestingly, clone 39 has shown generally high resistance to both biotic and abiotic threats (Fig. 5, Rousi et al. 1996b). The same clone also seems to be exceptionally resistant to ozone injury (clone 5-M, in P_ifikk6nen et al. 1993), whereas the generally herbivore susceptible clone 36 is similarly very susceptible to ozone damage (2-M, in P_i_ikk6nenet al. 1993). Environment May Sometimes Affect Birch Resistance Our previous experiments with various birch species (8 birch species) and origins have indicated, as expected, that growth environment (shading or fertilization) does not have an effect on low resistance species (slow or fast growing), such as B. papyrifera, pubescens, maximowicziana, schmidtii or grossa. In high resistance species, fertilization increased the palatability of two species, B. platyphylla and B. ermanii, but did not affect another, B. resinifera. Shading had no clear effect on any of the tested species (Rousi et al. 1996a). European and Japanese white birches are closely related (Dugle 1966). However, fertilization did not diminish the hare resistance of European white birch (Rousi et al. 1991, 1993), but lowered that of Japanese white birch (Rousi et al. 1996a). On the other hand, shading tends to increase hare palatability of European white birch (Figs. 6 and 7), but its effect on Japanese white birch is negligible (Rousi et al. 1996a). The effect of shading on European white birch is, however, strongly dependent on genotype and site fertility. There were no statistical differences in resistance among three families that we tested in low fertility soils (Fig. 6). But, if seedlings are grown in high fertility soil, resistance was significantly different in two out of three families (ANOVA, Bonferroni t-test, p
P FERTILIZATION E A L ( A 0 - no shade T 60 50 = 50% shade ( A B 50 I 40 L I 30 T y 20 10 I N 0 D E 0 50 0 50 0 50 X V 5845 x V 5848 V 5845 x V 5781 V 5845 x V 5846 Figure 7.--Hare resistance of three B. pendula F2 families in two shade treatments. Seedlings were grown in nursery peat and obtained optimum fi3rtilization (For details see Rousi et al. 1996a). Cafeteria experiments as in Figure 1. Mean + S.E. SUMMARY We conclude that prevailing defense ttleories, (e.g., Carbon-Nutrient Balance, Growth-Differentiation Balance, etc.) may not be fully applicable to birch. The reaction of birch species, origins, andgenotypes to variable environments produces variable resurts, which depend on plant genotype, environment:and the particular mechanism of resistance to each pest type. Consequently, generalized conclusions about birch resistance should be made with great caution. LITERATURE CITED BRYANT, J.R, CHAPIN, F.S. [[I and KLEIN, D.R. 1983. Carbon/nutrient balance of boreal plants in relation to vertebrate herbivory. Oikos 40:357-368. BRYANT, J.R, TAHVANAINEN, J., SULKINOJA, M., JULKUNEN-THTTO, R., REICHARDT R, and GREEN, T. 1989. Biogeographic evidence for the evolution of chemical defense against mammalian browsing by boreal birch and willow. The American Naturalist 134: 20-34. CHIBA, S. and NAGATA, Y. 1968. Resistance to field mice and mountain hares in Betula-species and hybrids. (1) Comparison between species. Qji Institute t\)r Forest Tree Improvement. Technical Note 68:20-24 (In Japanese). CttIBA, S. and NAGATA, Y. 1969. Resistance to field mice and mountain hares in Betula-species and hybrids. (2) Comparison between parental species and hybrids. Oji Institute for Forest Tree Improvement. Technical Note 88: 153- 155 (In Japanese). CHIBA, S., NAGATA, Y., and TOMAKL T 11982. Variation of vole resistance in Larix leptolepis and comparison with the hybrid L gmelini and L leptolepis. Oji Institute for Forest Tree Improvement. Technical Note 209:6-9 (In Japanese). 44
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Page 3 and 4: DYNAMICS OF FOREST HERBIVORY: QUEST
Page 5 and 6: TABLE OF CONTENTS Seeking The Rules
Page 7 and 8: 32. Companion planting of the nitro
Page 9 and 10: GRIMALSKY, ¥.I., Research Institut
Page 11 and 12: POUTTU, ANTTI, Finnish Forest Resea
Page 13 and 14: persist in mature leaves and affect
Page 15 and 16: THE SINK-SOURCE HYPOTHESIS: TYPE AN
Page 17 and 18: late season defoliation decreases t
Page 19 and 20: BLANCHE, C.A., LORIO, EL., Jr., SOM
Page 21 and 22: NIEMELA, R, TUOMI, J,, and LOJANDER
Page 24: induced reactions have been studied
Page 28 and 29: summer, but the other sawfly specie
Page 30 and 31: LARSSON, S., BJ(_RKMAN, C., and GRE
Page 32 and 33: STAND AND LANDSCAPE DIVERSITY AS A
Page 34 and 35: not related to resistance to the se
Page 36 and 37: In contrast to the situation in eas
Page 38 and 39: ROOT, R.B. 1973. Organization of a
Page 40 and 41: We tlhus neglect two potential sour
Page 42 and 43: c__ c__ 1T1 Ill a) a) 0 0 0 e 1 o e
Page 44 and 45: c_ >l.s S S S m N < DN DN < D c DN
Page 46 and 47: :hen k > 0 suggest that the cue or
Page 48 and 49: oth as protective weapons and as po
Page 50 and 51: DEFENSE THEORIES AND BIRCH RESISTAN
Page 52 and 53: P 45 A L 40 A T 35 A B 30 I L 25 I
Page 56 and 57: DUGLE, J.A. 1966. A taxonomic study
Page 58 and 59: accommodate findings related to mic
Page 60 and 61: Table l.--Mean ±S.E. area eaten by
Page 62 and 63: Sulfhydryl-Disulfide Mechanisms In
Page 64 and 65: Iraa prior experimental analysis of
Page 66 and 67: FRAZIER, J.L. and HEITZ, J.R. 1975.
Page 68 and 69: REICHARD, R 1993. From RNA to DNA,
Page 70 and 71: ; trees, and between and within ind
Page 72 and 73: Encapsulated objects are data struc
Page 74 and 75: _re3.--The Lignum model can be cont
Page 76 and 77: SEVERE DEFOLIATION Tree Fine Root +
Page 78 and 79: RUMBAUGH, J., BLAHA, M., PREMERLANI
Page 80 and 81: oo0 a b ol ¢' _achiman_i ()® 1,eA
Page 82 and 83: 1000 Conspicuous Defo i at ion 100
Page 84 and 85: 2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
Page 86 and 87: Qualitative Changes in [,eaves and
Page 88 and 89: t00 2 Yr 1 Yr o_ 80 Treatment Treat
Page 90 and 91: Table 2.--Body size of Q. purzctate
Page 92 and 93: 06 b AddedBSA < 04 _ 2rag -'=' i:3
Page 94 and 95: Severe Defol iat ion Site-A 1993 _
Page 96 and 97: KAMATA, N. and IGARASHI, M. 1995b.
Page 98 and 99: enclosure %r 20 post-diapausing sec
Page 100 and 101: 2,500 F 000 F NF NF b _7 a _;_ a _7
Page 102 and 103: 8O D 09 i 4,'.) F AS4L a Z___7 2o !
Page 104 and 105:
2,500 _ st2-st4 El st5 0 st6 [_ pup
Page 106 and 107:
96 SLANSKY, E, Jr. 1990. Insect nut
Page 108 and 109:
later, we selected one more floweri
Page 110 and 111:
Table 2.---Mean spruce budworm perf
Page 112 and 113:
Table 3..--Mean spruce budworm perf
Page 114 and 115:
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
Page 122 and 123:
THE BLACKMARGINED APHID AS A KEYSTO
Page 124 and 125:
Wood et al. (1987) report that M. c
Page 126 and 127:
MIZELL, R.E 1991. Pesticides and be
Page 128 and 129:
METHODS Study Site The study took p
Page 130 and 131:
the samples. Fertilization had subs
Page 132 and 133:
Table 2.--Percent nutrient content
Page 134 and 135:
Some tree responses to fertilizatio
Page 136 and 137:
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.
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A° ADDITIVE C. HYBRID SUSCEPTIBILI
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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
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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
Page 156 and 157:
ACKNOWLEDGEMENTS The authors sincer
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SINGH, D.R 1986_ Breeding for resis
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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
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-
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OGAWA, A., NAGATA, Y., and SUKENO,
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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 190 and 191:
Considering biochemical pathways, i
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
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1960, Kalkstein 1976). Increased su
Page 208 and 209:
RYAN, B.F., JOtNER, B.L., and RYAN,
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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
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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 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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