High foliar nitrogen concentration ([N]) is associated with high rates of photosynthesis and thus high tree productivity; however, at excessive [N], tree productivity is reduced. Reports of excessive [N] in the Douglas-fir forests of the Oregon Coast Range prompted this investigation of growth and needle physiological responses to increasing foliar N concentrations in 1-year-old Douglas-fir seedlings. After 1 year of N fertilization, total seedling biomass increased with each successive increase in N fertilizer concentration, except in the highest N fertilization treatment. Of the many physiological responses that were analyzed, only photosynthetic capacity (i.e., Vcmax), respiration rates and leaf specific conductance (KL) differed significantly between N treatments. Photosynthetic capacity showed a curvilinear relationship with foliar [N], reaching an apparent maximum rate when needle N concentrations exceeded about 12 mg g(-1). In vitro measurements of ribulose-1,5-bisphosphate carboxylase (Rubisco) activity suggested that photosynthetic capacity was best related to activated, not total, Rubisco content. Rubisco activation state declined as foliar [N] increased, and based on its significant correlation (r2= 0.63) with foliar Mn:Mg ratios, it may be related to Mn inactivation of Rubisco. Respiration rates increased linearly as foliar N concentration increased (r2= 0.84). The value of K(L) also increased as foliar [N] increased, reaching a maximum when foliar [N] exceeded about 10 mg g(-1). Changes in K(L) were unrelated to changes in leaf area or sapwood area because leaf area to sapwood area ratios remained constant. Cumulative effects of the observed physiological responses to N fertilization were analyzed by modeling annual net CO2 assimilation (Anet) based on treatment specific values of Vcmax, dark respiration (Rdark) and KL. Estimates of Anet were highly correlated with measured total seedling biomass (r2= 0.992), suggesting that long-term, cumulative effects of maximum Rubisco carboxylation, Rdark and KL responses to N fertilization may limit seedling production when foliar N exceeds about 13 mg g(-1) or is reduced to less than about 11 mg g(-1).
Summary
We investigated the nutritional dynamics of Phaeocryptopus gaeumannii and the impact of nitrogen (N) fertilization of Douglas‐fir (Pseudotsuga menziesii) on the production of P. gaeumannii fungal fruiting bodies. Emergence of P. gaeumannii fungal fruiting bodies (pseudothecia) in Douglas‐fir stomata has been directly linked to premature needle loss, a symptom of Swiss needle cast disease.
Douglas‐fir trees (10‐yr‐old) naturally infected with P. gaeumannii were treated with soil applications of N fertilizer isotopically enriched with 15N to increase foliar N and track the movement of N from the host to the fungus. Foliar N, free amino acids, percent of stomata occluded by pseudothecia, N isotope and carbon (C) isotope levels were assessed on treated and control trees.
Higher foliar N resulted in increased %N and %C in P. gaeumannii, as well as increased fungal fruiting and thus disease severity. Comparisons of δ15N levels between P. gaeumannii pseudothecia and associated needles indicated an increase in δ15N of needles and a simultaneous decline in δ15N of pseudothecia coupled with increased levels of foliar and fungal percentage N.
These findings confirm that P. gaeumannii responds to host nutrient status and that increased N availability inside Douglas‐fir needles is linked to increased severity of Swiss needle cast disease.
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