BY JARMO K. HOLOPAINEN, RISTO RIKALA, PIRJO KAINULAINEN AND JARI OKSANEN
New Phytologist 131, 521-532 (1995)
We tested how variable nitrogen availability affects the above and below ground growth of first-year Scots pine (Pinus sylvestris L.) seedlings and carbon allocation to defensive allelochemicals and storage. Levels of free amino acids were considered as indicators of nutritive quality. Suitability of seedlings for polyphagous Lygus rugulipennis Popp. (Heteroptera: Miridae) was tested with oviposition preference and nymphal growth experiments. At the end of the growing season, needle length increased while root biomass decreased with elevated N fertilization, but shoot length was not affected. Concentration of starch in needles and roots, representing carbon storage, was not significantly affected by the level of N fertilization, although there was a decreasing trend in the starch concentration of needles when nitrogen input increased. Nitrogen fertilization significantly increased the pool of total and non-essential amino acids in the shoots. Arginine, proline, and glutamine were most distinctively affected individual amino acids by elevated N. Of the carbon-based defence compounds, especially total resin acid concentrations in shoots were significantly reduced with elevated nitrogen in nine-weeks-old seedlings. Palustric acid and neoabietic acid were the most affected individual resin acids, while foliar monoterpenes were not influenced by N availability. Total phenolics in shoot and root showed variable response. The increasing effect of nitrogen on the oviposition rate of Lygus females was almost linear. Mean relative growth rate of the nymphs was significantly affected by the level of N fertilization but, the mortality of nymphs was high in all treatments. The results suggest that in nitrogen-rich environments the needle growth of small Scots pine seedlings is improved, but their susceptibility to insect attack is increased and they remain less defended as predicted by carbon/nutrient balance hypothesis. Total phenolics and resin acids, representing phenylalanine and mevalonic acid pathway, respectively, were both reduced with nitrogen availability. Together with the simultaneous increase of foliar free nitrogen in the form amino acids, the nutritive value of seedlings is ameliorated and this may explain susceptibility of nursery-grown, N fertilized seedlings to polyphagous Lygus bugs.
Key words: Fertilization, herbivory, Pinus sylvestris, plant chemistry, plant-insect interaction