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SEVERE DEFOLIATION Tree Fine Root + _ /_' / ........................ _ Carbohydrate Reserves Biomass - J_ Photosynthesis 14Y'+ Nutrient + Res erv es 1 + Tree ee Number / .,._.._ Carbohydrate Uptake Concentration Needle Size Capacity 'X_+ Concentration Needle Nutrient I + Nutrient | Needle C-Based Absorption , + Secondary Needle Food Quality _ Metabolites (No Induction) - Figure 6.--Dynamics of nutrients and carbohydrates in the Lignum model, with the effects of"defoliation. The present model, as it has explicit descriptions for foliage in each shoot, is capable of duplicating this kind of dynamics. However, we are still calibrating the model, and can not yet claim error-free predictions. DISCUSSION The present model was created for a testbench of ideas in many different disciplines. It should be reusable for diagnostic reasoning, studying the effects of herbivore attack, visualization of symptoms, research on growth and development, and modelling tree architecture. Although we are still working on the first version of the model, the experiences have been promising. Future versions will be built to incorporate simulation on parallel computer hardware, and zooming in and out of the tree. The color schemes that we use for illustrating the status of the functions of the tree do not produce realistic visualization of the symptoms as of now. We will study the color effects that follow from various injuries, and try to animate the suffering and death of trees. The model is a part of a larger system for integrated forest health management (Saarenmaa et al. 1994). It has been built compatible with object-oriented descriptions of forest insects and fungi. These have attributes that describe what parts of trees, represented as object classes, they attack and what kind of disturbances they cause in those parts. Matching OO trees with OO insects and other harmful agents will create new kind of ecological simulations and also be useful in diagnostic reasoning. SUMMARY A model of a tree which consists of thousands of small interconnected parts can potentially be useful for studying the effects of insect and disease attack on the whole system. We present such a model for Pinus sylvestris, which is based on object-oriented programming. The model consists of classes for shoots, buds, the stem, branch whorls, foliage, bark, phloem, sapwood, and heartwood. The model simulates primary production, respiration, growth, water flow, and starch flow, and has a geometry that can be monitored in 3-D color on computer screen. The effects of attacks by shoot boring bark beetles and defoliating insects are studied. 65
LITERATURE CITED AHONEN, J. and SAARENMAA, H. 1991. Model-based reasoning about natural ecosystems: An algorithm to reduce the computational burden associated with simulating multiple biotic agents, 193-200. In Computer Science for Environmental Protection. Sixth Symposium. Munich, Germany, December 4-6, 1991. Springer-Verlag, Heidelberg. BRATKO, I., MOZETIC, I., and LAVRAC, N. 1989. Kardio. A study in deep and qualitative knowledge for expert systems. 260 p. The MIT Press, Cambridge, Massachusetts. ERICSSON, A., HELLQVIST, B., LANGSTROM, B., LARSSON, S., and TENOW, O_ 1985. Effects on grow*h of simulated and induced shoot pruning by Tomicus piniperda as related to carbohydrate and nitrogen dynamics in scots pine. LAppl. Ecol. 22: 105-124. •FAGERSTROM, T., LARSSON, S., LOHM, U., and TENOW. O. 1977. Growth in scots pine (Pinus sitvestris L.): a hypothesis on response to Blastophagus piniperda L. (Col., Scolytidae) attacks. For. Ecol. and Manage. 1:273-281. HERMS, D.A. and MATTSON, W.J. 1992. The dilemma of plants: to grow or defend. The Quart. Rev. Biol. 67(3): 283- 335. KUKKOLA, M., ANNILA, E., TIMONEN, M., and VARAMA, M. 1994. The development and yield of the Scots pine stand repeatedly attacked by the pine shoot beetles. Manuscript. Finnish Forest Research Institute. KURTH, W. 1992. Morphological models of plant growth: Possibilities and ecological relevance. 14 p. ISEM's 8th International Conference on the State-of-the-Art in Ecological Modelling, Kiel, Germany, September 28 - October 2, 1992. LANGSTROM, B. and HELLQVIST, C. 1991. Effects of different pruning regimes on growth and sapwood area of Scots pine. For. Ecol. and Manage. 44: 239-254. LANGSTROM, B. and HELLQVIST, C. 1992. Height growth recovery and crown development in top-damaged Pinus sylvestris trees. Scandinavian J. For. Res. 7: 237-247. L,_NGSTROM, B., TENOW, O., ERICSSON, A., HELLQVIST, C., and LARSSON, S. 1990. Effects of shoot pruning on stem growth, needle biomass, and dynamics of carbohydrates and nitrogen in Scots pine as related to season and tree age. Can. J. For. Res. 20: 514-523. LINDENMEYER, A, 1968. Mathematical models for cellular interaction in development. Parts I and II. J. Theoret. Biol. 18: 280-315. MANDELBROT, B.B. 1977. The fractal geometry of nature. 468 p. W.H. Freeman and Co., New York. NIKINMAA, E. 1992. Analyses of the growth of Scots pine; matching structure with functions. Acta Forestalia Fennica 235, 68 p. PERTTUNEN, J., SIEVANEN, R., SALMINEN, H., VAKEVA, J., NIKINMAA, E., and SAARENMAA, H. (1994). i generic growth model for a tree based on simple structural units and object-oriented programming. 30 p. (Subn-_tted). PRUNSINKIEWICZ, E and HANAN, J. 1989. Lindenmayer systems, fractals, and plants. Lecture Notes in Biomathernat_ its 79, 120 p. de REFFYE, P., LECOUSTRE, R., EDELIN, C., and DINOUARD, P. 1989. Modelling plant growth and architecture, p. 237-246. In Cell to Cell Signalling: from Experiments to Theoretical Models. Academic Press, Oxford. 66
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DYNAMICS OF FOREST HERBIVORY: QUEST
Page 5 and 6:
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
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 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
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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
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Only recently have we conducted stu
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esistance to beetle attack. Another
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LORIO, EL., Jr'. and HODGES, J.D. 1
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EFFECTS OF ROOT INHABITING INSECT-F
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OCCURRENCE OF ROOT AND STEM SUBCORT
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]Predisposition of Root-infected Tr
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chemistry associated with root colo
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Implications to Natural Resource Ma
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KLEPZIG, K.D., RAFFA, K.F., and SMA
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RAFFA, K.F., PHILLIPS, T., and SALO
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METHODS The study was installed in
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FURNISS, M.M., LIVINGSTON, R.L., an
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METHODS In the spring and summer of
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The importance of a compound as a r
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WERNER, R.A. 1995. Toxicity and rep
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41 40 42 \ 4t 42 41 42 ",,,\ 42 40
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Table 1.--Summary data for the 26 p
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One weakness of using data from gen
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Lower Peninsula (populations 14-26)
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AUCLAIR, A.ND., WORREST, R.C., LACH
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KELLOMAKI, S., HANNINEN, H., and KO
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WARGO, RM. and HAACK, R.A. 1991. Un
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investigation. Sampling was done on
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Figure 2.--Changes in protein preci
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q) J 400 T J ! i i i i o control tr
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attacked trees than in control tree
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ACIDIC DEPOSITION, DROUGHT, AND INS
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Table 1.--Impact of acidic fog upon
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MENGEL, K., BREININGER, T., and LUT
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THE RESISTANCE OF SCOTCH PINE TO DE
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EXPERIMENTAL METHODS Experiments we
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C w Q. 1,200 1,000 o 800 C_ --- 600
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0 .................................
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FONTAtNE, R.G. 1985. Forty years of
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Host Defenses An important componen
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Functional Heterogeneity of Forest
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The Connectivity Index (CI). The CI
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FUNCTIONAL HETEROGENEITYANALYSIS :
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The Modified Hazard Map. The next l
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systems facilitate mapping of fores
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KOLASA, J. and ROLLO, C.D. 1991. In
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) i,i_i)i_)i_i_ U.S. Departme_]t of
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