Plant species are hypothesized to impact ecosystem nitrogen cycling in two distinctly different ways. First, differences in nitrogen use efficiency can lead to positive feedbacks on the rate of nitrogen cycling. Alternatively, plant species can also control the inputs and losses of nitrogen from ecosystems. Our current understanding of litter decomposition shows that most nitrogen present within litter is not released during decomposition but incorporated into soil organic matter. This nitrogen retention is caused by an increase in the relative nitrogen content in decomposing litter and a much lower carbon-to-nitrogen ratio of soil organic matter. The long time lag between plant litter formation and the actual release of nitrogen from the litter results in a bottleneck, which prevents feedbacks of plant quality differences on nitrogen cycling. Instead, rates of gross nitrogen mineralization, which are often an order of magnitude higher than net mineralization, indicate that nitrogen cycling within ecosystems is dominated by a microbial nitrogen loop. Nitrogen is released from the soil organic matter and incorporated into microbial biomass. Upon their death, the nitrogen is again incorporated into the soil organic matter. However, this microbial nitrogen loop is driven by plant-supplied carbon and provides a strong negative feedback through nitrogen cycling on plant productivity. Evidence supporting this hypothesis is strong for temperate grassland ecosystems. For other terrestrial ecosystems, such as forests, tropical and boreal regions, the data are much more limited. Thus, current evidence does not support the view that differences in the efficiency of plant nitrogen use lead to positive feedbacks. In contrast, soil microbes are the dominant factor structuring ecosystem nitrogen cycling. Soil microbes derive nitrogen from the decomposition of soil organic matter, but this microbial activity is driven by recent plant carbon inputs. Changes in plant carbon inputs, resulting from plant species shifts, lead to a negative feedback through microbial nitrogen immobilization. In contrast, there is abundant evidence that plant species impact nitrogen inputs and losses, such as: atmospheric deposition, fire-induced losses, nitrogen leaching, and nitrogen fixation, which is driven by carbon supply from plants to nitrogen fixers. Additionally, plants can influence the activity and composition of soil microbial communities, which has the potential to lead to differences in nitrification, denitrification and trace nitrogen gas losses. Plant species also impact herbivore behaviour and thereby have the potential to lead to animal-facilitated movement of nitrogen between ecosystems. Thus, current evidence supports the view that plant species can have large impacts on ecosystem nitrogen cycling. However, species impacts are not caused by differences in plant quantity and quality, but by plant species impacts on nitrogen inputs and losses.
