Nitrogen, parasites and plants : key interactions in boreal forest ecosystems

Abstract: In the work described in this thesis I studied how increases in nitrogen (N) inputs may affect plant community structure in boreal forest understorey vegetation. These phenomena were investigated in N fertilization experiments and along a national N deposition gradient. After five years of N additions, large changes in understorey vegetation composition were observed in the fertilization study. In plots that received 50 kg N ha'1 year"1 (N2), the abundance of the dominant species, Vaccinium myrtillus, decreased on average by 32 %. No decrease was observed in control plots during the same period. In contrast, the grass Deschampsia flexuosa responded positively to increased N input, being on average more than five times as abundant in the N2 treatments as in controls.Also an increase was seen in the incidence of disease caused by the parasitic fungus Valdensia heterodoxa on leaves of V. myrtillus following N additions. The parasite was on average nearly twice as abundant in N2 plots than in control plots. This could be explained by increased N concentrations in host plant tissue. Disease incidence also increased following experimental additions of glutamine to leaf surfaces of V. myrtillus, suggesting a causal connection between plant N concentration and performance of the fungus. The parasite also played a key role in the observed changes in understorey species composition. D. flexuosa was more abundant in patches in which V. myrtillus was severely affected by V heterodoxa. This suggests that V heterodoxa mediates the increased abundance of D. flexuosa following increased N additions. The fungus mediates changes in the composition of understorey vegetation mainly by increasing light availability via premature leaf loss of V. myrtillus.The incidence of disease due to the parasite was on average higher in large than in smaller N-treated plots, indicating that the response to N fertilization is spatially scale dependent. This shows that using small plot sizes in experiments that simulate changed environmental conditions may be problematic, as important interactions may be underestimated.Comparison of the occurrence of understorey species between regions with different rates of N deposition revealed that the occurrence of the two dwarf shrubs V. myrtillus and V. vitis- idaea was lower in regions with high N deposition compared to regions with low deposition. The opposite pattern was found for V heterodoxa. This is consistent with expectations from N fertilization experiments. For D. flexuosa no differences in occurrence were found between the different regions investigated.The effects on vegetation and mycorrhizal fungi observed following N additions were also found to be long lasting. Nine years after termination of the fertilization, no signs of recovery were detected, and nearly 50 years after termination characteristic signs of N fertilization were found among bryophytes and mycorrhizal fungi. This suggests that the time needed for re-establishment of the original biota following N-induced changes may be substantial.