View or print this publication - Northern Research Station - USDA ...

More documents

Recommendations

Info

cannot be derived from any other hypothesis: within the canopy, competition of resources among meristems puts a single branch with defoliation-weakened meristems into a poorer relative position if all other branches are intact than if they also are i weakened. Second, the S/S hypothesis provides a simple explanation for the multiannual "memory" of DIR (Haukioja el al. i 988): it might simply result from the time :needed to restore the vigor of meristems. Furthermore, the implications of the S/ S hypothesis also make it worthwhile to study whether plants have evolved mechanisms to prevent herbivores causing IA. For instance, it has been reported that developing current year foliage of Scots pine produces 13-keto-8(14)-podocarpen-18oic acid which deters pine sawflies and decreases use of the young current year foliage (Niemel_i et al. 1982). A strong deterrence of herbivores from such current year foliage may reduce the danger of induced amelioration. Perhaps the most important empirical message of the S/S hypothesis is a call for extreme caution in damage simulations. For instance, when triggering the DIR in the foliage of the mountain birch, even small variations in timing (perhaps a day or two) of defoliations may shift the treatment from sink leaves to source leaves. That may alter the response from IA to RIR or DIR, i.e., even the direction of the response. We doubt whether any study published so far has reported both the timing of defoliations in relation to tree phenology, and the exact defoliation practice, in such detail that it would allow the treatments to be precisely repeated. Actually, we are lacking a proper classification of damages to plants. Therefore, it is not astonishing that reports of plant responses to defoliations are notoriously variable (Lawton 1991). Although insects and plants may differ with reference to species, populations and even individuals, we regard the main problem the superficial knowledge of the system under study: we have to understand and to be able to record what we are doing with our own trees when we simulate herbivore damage. SUMMARY The debate about damage-induced changes in food plant quality has concentrated on their active vs incidental defensive roles. We show for birch and pine that the feedback consequences for herbivores vary from favorable to harmful depending on the treated plant part and on timing of the damage. Our results cannot be directly derived from the nutrient status of the tree. Instead, damage-induced alterations in meristematic sink strength provides a general explanation. In birch, damage to dominant sinks allows other parts to obtain a greater than normal share of resources which leads to their vigorous growth and better tbrage quality for herbivores. However, damage to young source leaves that are supplying local meristems for next year, weakens such meristems and reduces their success in competing for resources. This leads to less vigorous growth and poorer plant quality for herbivores. Damage to mature leaves presumably disturbs provisioning of the general resource pool of the tree but such effects, especially if they are monitored locally, may be hard to reveal. An emphasis on damage-induced effects on plant sinks and sources makes induced resistance, induced amelioration, as well as localization of responses more understandable. ACKNOWLEDGEMENTS We thank all the persons who have helped in the practical work on which this paper is based, as well as those who have helped in formulating the ideas and who have criticized earlier versions of the present paper (Sinikka Hanhim_iki, Seppo Neuvonen, Pekka Niemelfi, Matti Rousi, Kai Ruohom_.ki, Janne Suomela, John Vranjic, and Timo Vuorisalo). John also checked our English. Financially this study has been made possible by grants from the Academy of Finland, Maj and Tot Nessling Foundation, and the Lapland Forest Damage Project. LITERATURE CITED *GREN, G.I. and AXELSSON, B. 1985. Correlations between needle and shoot growth characteristics in Scots pine (Pinus sylvestris L.) stands. Forestry 58: 199-205. BENZ, G. 1974. Negative Riickkoppelung durch Raum- und Nahrungskonkurrenz sowie zyklische VerSnderung dr Nahrungsgrundlage als Regelprinzip in der Populationsdynamik des Grauen L_irchenwicklers, Zeiraphera diniana (Guende) (Lep., Tortricidae). Z. ang. Ent. 76: 196-228. BJC)RKMAN, C., LARSSON, S., and GREE R. 1991. Effects of nitrogen fertilization on pine needle chemistry and sawfly performance. Oecologia 86: 202-209. 7
BLANCHE, C.A., LORIO, EL., Jr., SOMMERS, R.A., HODGES, J.D., and NEBEKER, T.E. 1992. Seasonal cambial growth and development of loblolly pine: xylem formation, inner bark chemistry, resin ducts and resin flow. Forest. Ecol. Manage; 49:151-165. BRYANT, J.E, CHAPIN, F.S., III, and KLEIN, D.R. 1983. Carbon/nutrient balance of boreal plants in relation to vertebrate herbivory. Oikos 40: 357-368. BRYANT, J.E, KUROPAT, EJ., REICHARDT, EB., and CLAUSEN, T.E 1991. Controls over the allocation of resources by woody plants to chemical antiherbivore defense, p. 83-102. In Palo, R.T. and Robbins, C.T., yds. Plant Defenses Against Mammalian Herbivores. CRC Press, Boca Raton, Florida. BRYANT, J.E, REICHARDT, EB., CLAUSEN, T.E, and WERNER, R.A. 1993. Effects of mineral nutrition on delayed inducible resistance in Alaska paper birch. Ecology 74: 2072-2084. CLINE, M.G. 1991, Apical dominance. Bot. Rev. 57:318-358. DANELL, K. and HUSS-DANELL, K. 1985. Feeding by insects and hares on birches earlier affected by moose browsing. Oikos 44: 75_81. EDENIUS, L. 1993. Browsing by moose on Scots pine in relation to plant resource availability. Ecology 74:2261-2269. ERICSSON, A. 1978i Seasonal changes in translocation of 14Cfrom different age-classes of needles of 20-year-old Scots pine trees (Pinus sylvestris). Physiol. Plant. 43: 351-358. FARMER, E.E. and RYAN, C.A. 1990. Interplant communication: Airborne methyl jasmonate induces synthesis of proteinuse inhibitors in plant leaves. Proc. Natl. Acad. Sci USA 87: 7713-7716. HANHIM,g, KI, S. I989i Induced resistance in mountain birch: defence against leaf-chewing insect guild and herbivore competition: Oecologia 81: 242-248. HANHIMJkKI, S. and SENN, J. 1992. Sources of variation in rapidly inducible responses to leaf damage in the mountain birch-insect herbivore system. Oecologia 91:318-331. HARTLEY, S.E. and LAWTON, J.H. 1991. Biochemical aspects and significance of the rapidly induced accumulation of phenolics in birch foliage, p. 105-132. In Tallamy, D.W. and Raupp, M.J., yds. Phytochemical Induction By Herbivores. John Wiley, New York. HAUKIOJA, E. 198Z Inducible defences of white birch to a geometrid defoliator, Epirrita autumnata, p. 199-203. In Visser, J.H, and Minks, A.K., yds. Insect-Plant Relationships. PUDOC, Wageningen. HAUKIOJA, E. I991a! Cyclic fluctuations in density: interactions between a defoliator and its host tree. Acta Oecol. 12: 7-88 HAUKIOJA, E. 1991b_ The influence of grazing on the evolution, morphology and physiology of plants as modular organisms. Philos. Trans. R. Soc. Loud. B. 333: 241-247. HAUKIOJA, E. and NEUVONEN , S. 1985. Induced long-term resistance in birch foliage against defoliators: defensive or incidental? E_61ogy 66:1303-1308. HAUKIOJA, E. and NEUVONEN, S. 1987. Insect population dynamics and induction of plant resistance: the testing of hypotheses, p _11'432. In Barbosa, E and Schultz, J.C., yds. Insect Outbreaks. Academic Press, San Diego. HAUKIOJA, E., KAPIAINEN, K., NIEMELA, P., and TUOMI, J. 1983. Plant availability hypothesis and other explana- 8
Page 1 and 2: "
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: late season defoliation decreases t
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 54 and 55: However, hares showed strong prefer
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
Page 116 and 117:
Table 3.--Lymantria monacha prefere
Page 118 and 119:
500- 400- i 300 v ¢ m a 200- I00 L
Page 120 and 121:
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
Page 138 and 139:
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 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
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
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 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
show all

View or print this publication - Northern Research Station - USDA ...

Create successful ePaper yourself

Delete template?

Save as template?