Foliar free polyamine and inorganic ion content in relation to soil and soil solution chemistry in two fertilized forest stands at the Harvard Forest,Massachusetts

Polyamines (putrescine, spermidine, and spermine) are low molecular weight, open-chained, organic polycations which are found in all organisms and have been linked with stress responses in plants. The objectives of our study were to investigate the effects of chronic N additions to pine and hardwood stands at Harvard Forest, Petersham, MA on foliar polyamine and inorganic ion contents as well as soil and soil solution chemistry. Four treatment plots were established within each stand in 1988: control, low N (50 kg N ha^-1 yr^-1 as NH₄NO₃), low N + sulfur (74 kg S ha^-1 yr^-1 as Na₂SO₄), and high N (150 kg N ha^-1 yr^-1 as NH₄NO₃). All samples were analyzed for inorganic elements; foliage samples were also analyzed for polyamines and total N. In the pine stand putrescine and total N levels in the foliage were significantly higher for all N treatments as compared to the control plot. Total N content was positively correlated with polyamines in the needles (P 0.05). Both putrescine and N contents were also negatively correlated with most exchangeable cations and total elements in organic soil horizons and positively correlated with Ca and Mg in the soil solution (P 0.05). In the hardwood stand, putrescine and total N levels in the foliage were significantly higher for the high N treatment only as compared to the control plot. Here also, total foliar N content was positively correlated with polyamines (P 0.05). Unlike the case with the pine stand, in the hardwood stand foliar polyamines and N were significantly and negatively correlated with foliar total Ca, Mg, and Mn (P 0.05). Additional significant (P 0.05) relationships in hardwoods included: negative correlations between foliar polyamines and N content to exchangeable K and P and total P in the organic soil horizon; and positive correlations between foliar polyamines and N content to Mg in soil solution. With few exceptions, low N + S treatment had effects similar to the ones observed with low N alone for both stands. The changes observed in the pine stand for polyamine metabolism, N uptake, and element leaching from the soil into the soil solution in all treatment plots provide additional evidence that the pine stand is more nitrogen saturated than the hardwood stand. These results also indicate that the long-term addition of N to these stands has species specific and/or site specific effects that may in part be explained by the different land use histories of the two stands.