Modeling changes in red spruce carbon balance and allocation in response to interacting ozone and nutrient stresses.

The simulation model TREGRO was developed to analyze the response of red spruce saplings to multiple stresses, such as drought, nutrient deficiency, and exposure to pollutants. The model provides a method of identifying changes in structural and non-structural carbon resources in the tree that may become measurable only after many years of exposure. The model is based on the assumption that the ability of plants to take up and use carbon, water, and nutrients depends on the interrelationships in availability among the three resources. Consequently, the model simulates the simultaneous cycling of these resources. In the model, the tree is divided into the following compartments: a canopy of leaves grouped by age class, branches, stem, and coarse and fine roots in a number of soil horizons. In each of these compartments we track three carbon pools: living structure, dead structure or wood, and total non-structural carbohydrate. The model calculates the photosynthesis of an entire red spruce tree each hour as a function of ambient environmental conditions and the availability of light, water, and nutrients; the daily redistribution of carbon throughout the plant; and the loss of carbon by respiration and senescence. To accomplish this task, the model tracks the flow of carbon dioxide to the sites of fixation within the leaves, the availability of light in the canopy, water and nutrient resources in each of three soil horizons, and the amounts of these resources taken up by the tree. Soil and plant water potentials, photosynthesis, and leaf respiration are simulated on an hourly timestep; nutrient uptake, allocation and growth are computed on a daily timestep. Through a set of example simulations, we demonstrate how the model can be used to examine the mechanisms by which plants respond to stresses experienced alone and in combination. The model was used to predict the growth decrease and the shifting pattern of carbon allocation expected for an isolated tree exposed to ozone and decreased nutrient availability due to acidic deposition. Decreased nutrient availability resulted in decreased growth and preferential carbon allocation to roots, which helped to alleviate the nutrient stress. Ozone stress also resulted in decreased plant growth but had the opposite effect on allocation patterns, with most of the growth reduction occurring in roots. The effect of simultaneous ozone and nutrient stress on tree growth was less than the sum of the independent single stresses, contrary to our expectation. This modeling approach can aid in evaluating the long-term effect of stress on resource availability, the potential for gradual deterioration of tree health under long periods of stress, and imbalances in growth accompanying shifts in carbon allocation caused by stress.