Interactive effects of fertilization and throughfall exclusion on the physiological responses and whole-tree carbon uptake of mature loblolly pine

Few studies have examined the combined effects of nutrition and water exclusion on the canopy physiology of mature loblolly pine (Pinus taeda L.). Understanding the impacts of forest management on plantation productivity requires extensive research on the relationship between silvicultural treatments and environmental constraints to growth. We studied the physiological responses of 18-year-old loblolly pine trees exposed to a combination of fertilization (fertilizer or no fertilizer) and throughfall (normal throughfall or throughfall exclusion). Gas exchange variables were measured in the upper and lower crown between 0900 and 1700 h from May to November in 1999. Needle fall was collected to estimate foliage mass and leaf area. Summer drought and throughfall exclusion significantly decreased predawn xylem pressure potential. Needle-level photosynthesis, transpiration, and stomatal conductance declined during the drought and were significantly lower in the throughfall exclusion treatment. Throughfall exclusion also reduced annual foliage mass and daily whole-crown photosynthesis and transpiration. In the normal throughfall treatment, fertilization had no effect on needle-level physiology, but increased annual foliage mass and whole-crown photosynthesis by 26% and 41%, respectively. With the exclusion of throughfall, however, annual foliage mass and daily whole-crown photosynthesis exhibited little response to fertilization. We conclude that greater nutrient availability enhances the carbon uptake of mature loblolly pine trees by stimulating foliage production, but the positive effects of fertilization on leaf area and carbon fixation are limited by low water availability.