Abstract
In northern boreal forests ground level species occur which may have the capacity to suppress tree seedling regeneration substantially, particularly in late successional stages. We investigated, through an experiment involving sequential manipulations, the effects of three biotic components operating at the ground layer, i.e. aboveground effects of the ericaceous shrub Empetrum hermaphroditum and of the feathermoss species Pleurozium schreberi and the below-ground effects of extramatrical mycorrhizal hyphae, on the seedling establishment, growth and nutrient acquisition of Pinus sylvestris during two growing seasons. Germination and seedling establishment of P. sylvestris were enhanced by reduction of the above ground components of E. hermaphroditum and P. schreberi but were not influenced by temporary disruption of mycorrhizal hyphae. Seedling growth and nitrogen content were, however, significantly increased when hyphal connections were disrupted and above ground influence of P. schreberi and E. hermaphroditum were reduced. Higher shoot:root ratios were also found in seedlings when hyphal connections were disrupted. Seedlings in treatments where P. schreberi was left intact and above ground interference by E. hermaphroditum was reduced had unusually low shoot:root ratios (1.6) after two growing seasons and significantly lower total amounts of nutrients after one growing season than were initially present in the seeds, indicative of a net nutrient drain during the early establishment phase. The experiment could not explain the mechanisms behind this unexpected nutrient drain. Possible rhizosphere interactions immobilizing nutrients and causing seedling nutrient losses are discussed. From the results of this study and other indirect evidence we hypothesize that the three biotic components we investigated could act together to immobilize nutrients and to inhibit tree seedling regeneration and growth in late successional stages of boreal forests.
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