Growth response of Douglas-fir seedlings to nitrogen fertilization: importance of Rubisco activation state and respiration rates

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).