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esistance to beetle attack. Another study is in progress, once again with juvenile pines, in a plantation subjected to thinning<br />

and fertilization. We will be continuing w<strong>or</strong>k in that direction. At <strong>this</strong> time preliminary results indicate that the sh<strong>or</strong>t-term<br />

effects of thinning and fertilization are to enhance carbon partitioning to growth and to reduce carbon committed to secondary<br />

metabolism; e.g., resin synthesis.<br />

SUMMARY<br />

There are a number of ways to approach the problem of assessing the effects of environmental conditions, such as<br />

water regime, on tree physiological responses and resistance to bark beetle attack. It helps to keep in mind that environmental<br />

fact<strong>or</strong>s operate through physiological processes (Fig. 1, and Kramer 1986), and that there are concepts, such as plant<br />

growth-differentiation balance (Loomis 1932) and carbon-nutrient balance (Bryant et al. 1983; Tuomi 1992; Tuomi et al.<br />

1988, 1990), that can provide bases f<strong>or</strong> f<strong>or</strong>ming testable hypotheses. It is especially imp<strong>or</strong>tant to know as much as possible<br />

about the host tree and the specific bark beetle of interest. Observational studies are particularly imp<strong>or</strong>tant because they can<br />

provide a baseline f<strong>or</strong> interpreting the results of manipulative studies. Whenever feasible, it is especially helpful in manipulative<br />

studies to characterize environmental conditions bef<strong>or</strong>e and during a study. Because physiological changes that occur<br />

during ontogeny of trees can alter their susceptibility to herbiv<strong>or</strong>es, knowledge of tree stage of growth and development can<br />

be especially helpful.<br />

ACKNOWLEDGMENTS<br />

Cooperative Agreements (19-86-036 and 19-87-20) between the Mississippi Agricultural and F<strong>or</strong>estry Experiment<br />

<strong>Station</strong> and the <strong>USDA</strong> F<strong>or</strong>est Service, Southern F<strong>or</strong>est Experiment <strong>Station</strong>, contributed in part to results rep<strong>or</strong>ted here.<br />

' LITERATURE CITED<br />

BRYANT, J.P., CHAPIN, F.S., III, and KLEIN, D.R. 1983. Carbon/nutrient balance of b<strong>or</strong>eal plants in relation to vertebrate<br />

herbiv<strong>or</strong>y. Oikos 40: 357-368.<br />

CHAPIN, ES., III, BLOOM, A.J., FIELD, C.B., and WARING, R.H. 1987. Plant responses to multiple environmental<br />

fact<strong>or</strong>s. BioScience 37: 49-57.<br />

CHOU, C.K.S. 1982. Susceptibility of Pinus radiata seedlings to infection by Diplodia pinea as affected by pre-inoculation<br />

conditions. New Zealand J. F<strong>or</strong>. Sci. 438-441:<br />

COLEY, P.D., BRYANT, J.P., and CHAPIN, F.S., III. 1985. Resource availability and plant antiherbiv<strong>or</strong>e defense. Science<br />

230: 895-899.<br />

DUNN, J.P. and LORIO, P.L., Jr. 1992. Effects of bark girdling on carbohydrate supply and resistance of loblolly pine to<br />

southern pine beetle (Dendroctonusfrontalis Zimm.) attack. F<strong>or</strong>. Ecol. and Manage. 50:317-330.<br />

DUNN, J.P. and LORIO, P.L., Jr. 1993. Modified water regimes affect photosynthesis, xylem water potential, cambial<br />

growth, and resistance of juvenile Pinus taeda L. to Dendroctonusfrontalis (Coleoptera: Scolytidae). Environ.<br />

Entomol. 22(5): 948-957.<br />

GORDON, J.C. and SMITH, W.H. 1987. Tree roots and microbes: a high pri<strong>or</strong>ity f<strong>or</strong> high technology, p. 423-432. In<br />

Boersma, L.L., et al., eds. Future Developments in Soil Science <strong>Research</strong>. Soil Science Society of America, Madison,<br />

WI.<br />

HEDDEN, R.L. and LORIO, RL., Jr. 1985. Rating stand susceptibility to southern pine beetle attack on national f<strong>or</strong>ests in<br />

the Gulf Coastal Plain. Res. Pap. SO-221. Asheville, NC: U.S. Department of Agriculture, F<strong>or</strong>est Service, Southern<br />

<strong>Research</strong> <strong>Station</strong>. 5 p.<br />

207

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