Effects of competition on the foliar chemistry of young Douglas-fir in monoculture and mixed stands with young red alder
The effects of Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) and red alder (Alnusrubra Bong.) densities on foliar chemistry of 4-year-old Douglas-fir were examined in experimentally manipulated stands. The density of each species was varied systematically in a two-way density matrix composed of five monoculture densities and 25 mixtures of all possible pairwise combinations of monoculture densities. Foliage was sampled in July of the second growing season and analysed for concentrations of nitrogen (N), phosphorus (P), starch, sugars, total nonstructural carbohydrates, and free amino acids. Response surfaces were generated, and regression analyses quantified the effects of each species' density on biochemistry. Increasing the overstory alder density resulted in increased N, free amino acid, and P concentrations in the Douglas-fir foliage, and reduced starch and total nonstructural carbohydrate concentrations. Effects of Douglas-fir density were variable and dependent on the red alder density. Trends in biochemistry correlated well with soil moisture, light, and growth rates. Conditions favoring growth were associated with reduced concentrations of N and P and elevated levels of starch and nonstructural carbohydrates. The results strongly suggest that examination of foliar chemistry should be coupled with growth, physiology, and resources if biochemical constituents are to be used for testing tree vigor.