USDA Forest Service

Pacific Southwest Research Station


Pacific Southwest Research Station
800 Buchanan Street
West Annex Building
Albany, CA 94710-0011

(510) 559-6300

United States Department of Agriculture Forest Service. USDA logo which links to the department's national site. Forest Service logo which links to the agency's national site.

Programs and Projects


Air Pollution and Global Change Impacts on Western Forest Ecosystems

Photo of Nancy Grulke

Nancy Grulke
Research Plant Physiologist
Project Leader, Forest Health and Disturbance
Employee E-mail Address Image

Pacific Southwest Research Station
Forest Fire Laboratory
4955 Canyon Crest Drive
Riverside, CA 92507

Ph: (951) 680-1500
Fx: (951) 680-1501

Research Interests/Duties

Research Assignment: To develop (a) an understanding of whole tree responses to environmental stressors and growth enhancers, (b) empirical models to predict tree responses, and (c) a scientific basis for remote detection of tree stress.

Research Interests: Ecophysiological and life historical characteristics of plants; Whole plant and ecosystem response to environmental stressors (ozone exposure, excess nitrogen deposition, drought) and growth enhancers (CO2 enrichment; low to moderate nitrogen deposition); Empirical modeling of physiological processes to single and multiple environmental influences.

Research Experience: Plant community and demography; Ecophysiological and life history research experience with grasses, forbs, shrubs, and all age classes of coniferous, evergreen broadleaf, and deciduous trees; Field research experience in the polar desert (Canada), arctic tundra (Alaska), boreal forest (Alaska, Maine), subalpine forests (Colorado), temperate coniferous forests (California, North Carolina, Italy), mediterranean oak woodlands (California, Italy), temperate beech forest (Bavaria), and coastal mangroves (Bahamas).

Research Approach: Basic science approach used in solving applied problems; Known responses to single environmental stressors as determined in exposure chambers are used to help interpret the interactive effect of multiple stressors in woodlands and forests; Tree morphological and physiological attributes are described in natural forest stands along known environmental gradients.

Selected Publications

Aberrant stomatal behavior:
Grulke NE, Neufeld HS, Davison AW, Chappelka A 2007 Stomatal behavior of O3-sensitive and -tolerant cutleaf coneflower (Rudbeckia laciniata var. digitata) Great Smoky Mountain National Park. New Phytologist 173:100-109.

Grulke NE, Paoletti E, Heath RL 2007 Chronic vs. short term acute O3 exposure effects on nocturnal transpiration in two Californian oaks. TheScientificWorldJournal (2007) 7(S1):134-140. DOI 10.1100/tsw.20007.33

Grulke, N.E., Paoletti, E., Heath, R.A. 2007 Comparison of calculated and direct measurements of foliar O3 uptake in crop and native tree species. Environmental Pollution 146:640-647.

Grulke NE, Paoletti E 2005 New system to deliver desired O3 concentrations in leaf-level gas exchange measurements: results for Holm oak near a CO2 spring. Phyton (Austria) 45:21-31.

Grulke NE, Dobrowolski W, Mingus P, Fenn ME 2005 California black oak response to N-amendment at an N-saturated site. Environmental Pollution 137:536-545.

Paoletti E, Grulke NE 2005 Does living in elevated CO2 ameliorate tree response to ozone? A review on stomatal responses. Environmental Pollution 137:483-493.

Grulke NE, Alonso R, Nguyen T, Cascio C, Dobrowolski W 2004 Stomata open at night: implications for pollutant uptake in ponderosa pine. Tree Physiology 24:1001-1010.

Grulke NE 2000 A statistical model for stomatal behavior under fluctuating light: the role of short-, medium- and long-term elevated ozone exposure. Pg 35, Proceedings, 91st Annual Meeting, IUFRO, Division 7.04, D Karnosky (tech coor), Symposium held May 26-30, 2000. Michigan Technological University, Houghton, MI.

Ecophysiological field studies of mediterranean-type forest species:
Staszak J, Grulke NE, Prus-Glowacki W 2007 Air pollution-driven genetic change in yellow pine in Sequoia National Park. Environmental Pollution 149:366-375.

Staszak J, Grulke NE, Prus-Glowacki W 2004 Genetic differences of Pinus ponderosa trees tolerant and sensitive to O3 exposure. Water, Air, Soil Pollution 153:3-14.

Grulke NE, Johnson R, Jones D, Monschein S, Nikolova P, Tausz M 2003 Variation in morphological and biochemical O3 injury attributes of Jeffrey pine within canopies and between microsites. Tree Physiology 23:923-929.

Grulke NE, Johnson R, Esperanza A, Jones D, Nguyen T, Posch S, Tausz M 2003 Canopy transpiration of Jeffrey pine in mesic and xeric microsites: O3 uptake and injury response. TREES 17:292-298.

Grulke NE, Preisler H, Rose C, Kirsch J, Balduman L 2002 Evaluating the role of drought stress on ozone uptake in ponderosa pine. New Phytologist 154: 154:621-632.

Grulke NE, Preisler H 2002 A statistical approach to estimate ozone uptake of ponderosa pine in a mediterranean climate. Environmental Pollution 119:163-175.

Grulke NE, Andersen CP, Hogsett WE 2001 Seasonal changes in carbohydrate pools of ponderosa pine in stands under differing environmental stress. Tree Physiology 21:173-184.

Grulke NE, Retzlaff WA 2001 Changes in physiological attributes from seedlings to mature ponderosa pine. Tree Physiology 21:275-286.

Retzlaff WA, Arthur MA, Grulke NE, Weinstein DA, Gollands B, Grulke NE 2000 Use of a single-tree simulation model to predict the effects of ozone and drought on a white fir tree. Tree Physiology 20:195-202.

Grulke NE, Balduman L 1999 Deciduous conifers: high nitrogen deposition and ozone exposure effects on ponderosa pine. Water, Soil, Air Pollution 116:235-248.

Grulke NE, Andersen CP, Fenn MP, Miller PR 1998 Ozone exposure and N deposition reduces root biomass in ponderosa pine across the San Bernardino Mountains, California. Environmental Pollution 103:63-73.

Grulke NE 1998 The role of early, late, and whole season ozone exposure in photosynthetic decline of ponderosa pine. Proceedings, Air Waste Management Assoc., Paper No. 98-TA37.05. Pittsburgh, PA: Air and Waste Management Assoc.

Grulke NE, Lee EH 1997 Assessing ozone-induced foliar injury in ponderosa pine. Canadian Journal of Forest Research 27:1658-1668.

Grulke NE, Miller PR, Scioli D 1996 Response of giant sequoia canopy foliage to experimentally manipulated levels of atmospheric ozone. Tree Physiology 16:575-581.

Grulke NE, Miller PR 1994 Change in physiological characteristics from seedlings to mature trees in giant sequoia: implications for response to current and future levels of atmospheric pollutants. Tree Physiology 14:659-668.

Chamber exposure studies:
Handley T, Grulke NE 2008 Interactive effects of O3 exposure on California black oak (Quercus kelloggii Newb.) seedlings with and without nitrogen amendment. Environmental Pollution 156:53-60.

Grulke NE, Miller PR, Leininger TD 1999 Effect of ozone exposure on seasonal gas exchange of five western conifers. USDA Forest Service, Albany, CA: PSW-GTR-165:229-238.

Grulke NE, Miller PR, Scioli D 1996 Response of giant sequoia canopy foliage to experimentally manipulated levels of atmospheric ozone. Tree Physiology 16:575-581.

Grulke NE, Hom JL, Roberts SW 1993 Elevated CO2 response of 2 full-sib families of ponderosa pine. Tree Physiology 12:391-401.

Grulke NE, Miller PR, Wilborn RD, Hahn S 1989 Photosynthetic response of giant sequoia seedlings and rooted branchlets of mature foliage to ozone fumigation. Pgs 429-442, In: Olson RK, Lefohn AS (eds) Effects of air pollution on western forests. Trans. Of the Air Pollution Control Association, No. 16, Pittsburgh, PA: Air and Waste Management Assoc.

Arctic research:
Grulke NE 1995 Distribution of Phippsia algida and autosuccession in the polar semi-desert. Arctic Alpine Research 27:172-179.

Hahn SC, Oberbauer SF, Gebauer R, Grulke NE, Lange OL, Tenhunen JD 1996 Aboveground carbon and nutrient pools in the Imnaviat Creek Wilderness. Pgs 109-128, In: Reynolds JF, Tenhunen JD (eds.) Landscape Function and Disturbance in Arctic Tundra. Ecological Studies 120, Berlin: Springer Verlag.

Oechel WC, Cowles S, Grulke NE, Hastings SJ, Lawrence B, Prudhomme T, Riechers G, Vourlitis G 1994 In situ acclimation of net CO2 flux to elevated CO2: limits to CO2 fertilization in a natural ecosystem. Nature 371:500-505.

Oechel WC, Jenkins M, Hastings SJ, Riechers GH, Grulke NE, Vourlitis G 1993 CO2 release to the atmosphere from arctic tundra ecosystems: possible positive feedbacks to climate change. Nature 361:520-523.

Oechel WC, Riechers GH, Lawrence WT, Prudhomme TI, Grulke NE 1992 CO2LT, a closed null balance system for long term in situ ecosystem manipulation and measurement of CO2 level, CO2 flux, and temperature. Functional Ecology 6:86-100.

Grulke NE, Riechers GH, Oechel WC, Hjelm U, and Jaeger C 1990 Carbon balance of tussock tundra under ambient and elevated atmospheric CO2. Oecologia 83:485-494.

Grulke NE, Bliss LC 1988 Comparative life history characteristics of two High Arctic grasses, N.W.T. Ecology 69:131-143.

Grulke NE Bliss LC 1985 Growth forms, carbon allocation, and reproductive strategies of high arctic saxifrages. Arctic Alpine Research 17:241-250.

Book Chapters:
Grulke NE, Paine T, Minnich R, Chavez D, Riggan P, Dunn A 2009 Air pollution increases forest susceptibility to wildfire. Pgs 365-403, In: Bytnerowicz, A., Arbaugh, M., Riebau, A, Andersen, C. eds. Wildland Fires and Air Pollution. Developments in Environmental Science, Vol. 8, The Hague, Netherlands: Elsevier Publishers.

Grulke NE 2003 Physiological basis of ozone injury assessment in Sierra Nevadan conifers. In: Bytnerowicz A, Arbaugh M, Alonso R. Assessment of ozone distribution and its effects on Sierra Nevada ecosystems. Developments in Environmental Science 2:55-81. Elsevier Publishers, The Hague, Netherlands.

Grulke NE 1999 Physiological responses of ponderosa pine to gradients of environmental stressors. In: Miller PR, McBride J (eds) Oxidant Air Pollution Impacts in the Montane Forests of Southern California: The San Bernardino Mountain Case Study. Ecological Studies 134:126-163. NY:Springer-Verlag.

Last Modified: Mar 28, 2013 02:54:58 PM