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One weakness of using data from genetic tests plantings to study the impact of climatic warming, however, is that other variables besides temperature change when moving seed from its site of origin to the test location. Photoperiod, for example, is one important variable for which many tree species, including paper birch, are genetically adapted (Vaartaja 1959, Wright 1976, Kozlowski et al. 1991, Farmer 1993). In the northern hemisphere, typically, northern sources tend to flush earlier and terminate shoot growth earlier when moved southward compared with more local sources (Wright 1976, Kozlowski et al. 1991). In the present study, birch seed was collected between Lat. 43.3°N and 47.4°N, and then planted at 42.0°N. On the longest day of the year there is about 15.2 hours of light at 42.0°N, 15.5 hours at 43.3°N, and 16.0 hours at 47.4°N. Similarly, on the shortest day of the year, they experience about 9.1 hours of light at 42.0°N, 9.0 hours at 43.3°N, and 8.4 hours at 47.4°N. As a means of minimizing any influence of photoperiod, linear regression analyses were also conducted on two subsets of the original 26 populations (collection sites) in which the latitudinal spread among the seed sources was further reduced. In the first case, only the 13populations from the Lower Peninsula of Michigan were tested (populations 14- 26, Fig. 2, Table 1), representing collections within a 2° span of latitude and a 5°F (2.8°C) span in annual mean temperature. In the second case, 12 birch populations were tested (populations 4-7, 10, 12-18; see Fig. 2, Table 1), representing collections within a 1.5° span in latitude and a 4°F (2.2°C) span in annual mean temperature. Analyses were conducted using SAS and SigmaStat. Height and borer data were analyzed on an individual tree basis, whereas tree survival was calculated on a family basis (n = 218), using only those birch families that had at least one tree surviving through the second growing season (1977). Each tree was assigned to one of four "treatments" based on the average annual mean temperature of its origin. The four groupings were isotherms 40 ° and 41°F, 42 ° and 43°F, 44 ° and 45°F, and 46 oand 47°F (Table 2). The climatic differential term assigned to each grouping was CD1 ° (46-47°F), CD2°(44°-45°F), CD3 ° (42°-43°F), and CD4 ° (40°-41°F; Table 2). These terms, CDI°-CD4 °, reflect the average climatic differential in °C fbr each treatment group, e.g., for CD1 ° the midpoint climatic differential value is actually 1.I°C and represents the average degree of climatic warming that the trees in isotherms 460F and 47°F were theoretically experiencing (Table 2). Analysis of variance (ANOVA) was used to test for differences among treatments; ap level of 0.05 was used for significance. Tree height and survival data were analyzed using the GLM procedure of SAS, followed by the Student-Newman-Keuls means separation test. The arcsin square root transformation was performed on the survival data prior to analysis. Borer data were analyzed using the Kruskal-Wallis nonparametric test of SigmaStat, which performs an ANOVA on ranks; Dunn's test was used to make multiple comparisons among treatment means. RESULTS Paper birch growth, survival, and susceptibility to bronze birch borer attack were significantly affected by the climatic differential between the origin of the seed source and the plantation site (Fig. 3). Overall, height growth decreased as the climatic differential increased (Fig. 3A). Differences in height growth were already evident by the end of the second growing season in 1977 (p < 0.0001; Fig. 3A). By the end of the sixth growing season (1985), CDI ° trees were the tallest, with tree height becoming successively shorter for the CD2 °, CD3 °, and CD4 ° trees (p < 0.0001 ; Fig. 3A). As the trees aged, tree survival tended to decrease as the climatic differential increased (Fig. 3B). From year 2 (1977) to year 8 (1983) post-establishment, little tree mortality occurred, but thereafter tree mortality accelerated, especially among the trees experiencing the greatest climatic differential (CD4°; Fig. 3B). Treatment survival values first became significantly different in year 12 post-establishment (1987; p < 0.006), when survival for the CD4 ° trees was lower than that for trees from the other three groups. A similar pattern of tree survival continued into years 15 (1990; p < 0.001) and 16 (1991; p < 0.0008) post-establishment. As the trees became older, tree susceptibility to the bronze birch borer increased with increasing climatic differential (Fig. 3C). Of the 146 paper birch trees that died between 1985 and 1991, 143 (98%) of them had signs of borer attack along their lower trunk. A similar pattern emerged during each of the 3 years when borer damage was assessed (1985, 1987, 1990; p < 0.001 for each), i.e., CD4 ° trees experienced the highest levels of borer attack, while CDI ° trees experienced the lowest levels (Fig. 3C). As would be expected from the above discussion, the occurrence of borer-related bark ridges and exit holes was more common on CD4 ° trees than on CD 1° trees (Table 2). Regarding the two subsets of birch populations in which the latitudinal spread was reduced, paper birch growth, survival, and susceptibility to bronze birch borer were all significantly affected by the climatic differential between the origin of the seed source and the plantation site. In the first case, linear regression analysis of the 13 populations from Michigan's 239
- A 7 A _" --l-- 6 5 C c_ -,.-- CD1 o "1- U_I 3 _ CD2 ° I'-- r'r" i"Ti 2 4 jc _ CD3 o 1 I_B +- CU4 ° __1 ! r 1 _ _ _ .... f i' _ ...... _ T ' i ' E 1 2 4 6 8 10 12 14 16 100 : ........... A A 0_ _ 70 ",_ ",,,_AA.. ,,u4,6o ¢7" B B !--'- 50 40 , ," , .......... 2 4 6 8 10 12 14 16 "_0_"/ , .... .., ...., ..,B , ...., , ., , , 2 4 6 8 10 12 14 16 TREE AGE (Years) Figure 3.kPaper birch mean height growth (A), mean survival (B), and mean susceptibility to borer attack (C) over time (years post-establishment) in a provenance plantation in southern Michigan. Survival was based on only those trees that lived for the first two years. The 0-3 borer rating scale is presented in the Methods section. Trees are divided into four treatment groups based on the climatic differential (CD) between the average annual mean temperature of the origin of each seed source and the annual mean temperature of the plantation site. On average, trees assigned to treatment CD 1° had a 1.1°C difference between the origin and the test site, CD2 ° a 2.2°C difference, CD3 ° a 3.3°C difference, and CD4 ° a 4.4*C difference (see Table 2). For each parameter, means followed by the same letter within a column (i.e., year) are not significantly different (p
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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
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
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Page 206 and 207: 1960, Kalkstein 1976). Increased su
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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 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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