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Identifying the thresholds for the positive responses of total net primary productivity (NPP) to nitrogen (N) enrichment is an essential prerequisite for predicting the benefits of N deposition on ecosystem carbon sequestration. However, the responses of below-ground NPP (BNPP) to N enrichment are unknown in many ecosystems, which limits our ability to understand the carbon cycling under the scenario of increasing N availability. We examined the changes in above-ground NPP (ANPP), BNPP, and NPP of a temperate meadow steppe across a wide-ranging N addition gradient (0, 2, 5, 10, 20, and 50 g N m -2 year -1 ) during 5 years. Both ANPP and NPP increased nonlinearly with N addition rates. The N saturation threshold for ANPP (T A ) and NPP (T N ) was at the rate of 13.11 and 6.70 g N m -2 year -1 , respectively. BNPP decreased with increasing N addition when N addition rates ˃5 g N m -2 year -1 , resulting in much lower T N than T A . Soil N enrichment played a key role in driving the negative impacts of high N addition rates on BNPP, and consequently on the earlier occurrence of N saturation threshold for NPP. Our results highlight the negative effects of soil N enrichment on NPP in natural grasslands super-saturated with N. Furthermore, by considering ANPP and BNPP simultaneously, our results indicate that previous findings from above-ground might have over-estimated the positive effects of N deposition on primary productivity.

Temporal stability of net primary productivity (NPP) is important for predicting the reliable provisioning of ecosystem services under global changes. Although nitrogen (N) addition is known to affect the temporal stability of aboveground net primary productivity (ANPP), it is unclear how it impacts that of belowground net primary productivity (BNPP) and NPP, and whether such effects are scale dependent. Here, using experimental N addition in a grassland, we found different responses of ANPP and BNPP stability to N addition at the local scale and that these responses propagated to the larger spatial scale. That is, N addition significantly decreased the stability of ANPP but did not affect the stability of BNPP and NPP at the two scales investigated. Additionally, spatial asynchrony of both ANPP and BNPP among communities provided greater stability at the larger scale and was not affected by N addition. Our findings challenge the traditional view that N addition would reduce ecosystem stability based on results from aboveground dynamics, thus highlighting the importance of viewing ecosystem stability from a whole system perspective.

Most mistletoes are xylem-tapping hemiparasites, which derive their resources from the host's xylem solution. Thus, they affect the host's water relations and resource balance. To understand the physiological mechanisms underlying the mistletoe-host relationship, we experimentally removed Viscum album ssp. austriacum (Wiesb.) Vollmann from adult Pinus sylvestris L. host trees growing in a Swiss dry valley. We analyzed the effects of mistletoe removal over time on host tree growth and on concentrations of nonstructural carbohydrates (NSC) and nitrogen (N) in needles, fine roots and sapwood. In addition, we assessed the δ13C and δ18O in host tree rings. After mistletoe removal, δ13C did not change in newly produced tree rings compared with tree rings in control trees (still infected with mistletoe), but δ18O values increased. This pattern might be interpreted as a decrease in assimilation (A) and stomatal conductance (gs), but in our study, it most likely points to an inadequacy of the dual isotope approach. Instead, we interpret the unchanged δ13C in tree rings upon mistletoe removal as a balanced increase in A and gs that resulted in a constant intrinsic water use efficiency (defined as A/gs). Needle area-based concentrations of N, soluble sugars and NSC, as well as needle length, single needle area, tree ring width and shoot growth, were significantly higher in trees from which mistletoe was removed than in control trees. This finding suggests that mistletoe removal results in increased N availability and carbon gain, which in turn leads to increased growth rates of the hosts. Hence, in areas where mistletoe is common and the population is large, mistletoe management (e.g., removal) may be needed to improve the host vigor, growth rate and productivity, especially for relatively small trees and crop trees in xeric growth conditions.