Understanding and quantification of phosphorus (P) fluxes are key requirements for predictions of future forest ecosystems changes as well as for transferring lessons learned from natural ecosystems to croplands and plantations. This review summarizes and evaluates the recent knowledge on mechanisms, magnitude, and relevance by which dissolved and colloidal inorganic and organic P forms can be translocated within or exported from forest ecosystems. Attention is paid to hydrological pathways of P losses at the soil profile and landscape scales, and the subsequent influence of P on aquatic ecosystems. New (unpublished) data from the German Priority Program 1685 ''Ecosystem Nutrition: Forest Strategies for limited Phosphorus Resources'' were added to provide up-to-date flux-based information. Nitrogen (N) additions increase the release of water-transportable P forms. Most P found in percolates and pore waters belongs to the so-called dissolved organic P (DOP) fractions, rich in orthophosphate-monoesters and also containing some orthophosphate-diesters. Total solution P concentrations range from ca. 1 to 400 μg P L -1 , with large variations among forest stands. Recent sophisticated analyses revealed that large portions of the DOP in forest stream water can comprise natural nanoparticles and fine colloids which under extreme conditions may account for 40-100% of the P losses. Their translocation within preferential flow passes may be rapid, mediated by storm events. The potential total P loss through leaching into subsoils and with streams was found to be less than 50 mg P m -2 a -1 , suggesting effects on ecosystems at centennial to millennium scale. All current data are based on selected snapshots only. Quantitative measurements of P fluxes in temperate forest systems are nearly absent in the literature, probably due to main research focus on the C and N cycles. Therefore, we lack complete ecosystem-based assessments of dissolved and colloidal P fluxes within and from temperate forest systems. This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. Forest phosphorus cycle during ecosystem developmentForests are complex biogeochemical systems in which nutrient cycles readily change and become re-adjusted upon interactions with biotic and abiotic controls over diurnal, annual, decadal, centennial, and longer timescales (Hedin et al., 2003). Phosphorus (P) is an essential element for all living organisms. Modern agriculture avoids P limitation of primary production by continuous application of fertilizers, while forest ecosystems have developed efficient strategies for adapting to low P supply (Elser et al., 2007;Ilg et al., 2009;Rennenberg and Schmidt, 2010;Hinsinger et al., 2011). Increasing production of forests biomass in response to high atmospheric nitrogen (N) input and climate c...

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Natural fine colloids and nanoparticles have the potential to encapsulate and bind nutrients. Their size and composition is therefore relevant to understand the transport of essential nutrients like phosphorus in an aquatic ecosystem. The aim of this study was to characterize fine colloidal and nanoparticulate bound P of distinct hydromorphological areas in stream water from a forested test site in a small headwater catchment. Asymmetric flow field flow fractionation coupled online to inductively coupled plasma mass spectrometry was applied for size‐resolved detection of P, Fe, and Al in the fractions. Online P detection was a challenge due to the low concentrations (in this study down to 0.1 μg/L) in many natural waters. Additionally, the “dissolved” organic matter (DOM) content was derived from the online UV signal. The colloidal P occurred in two size fractions (2–20 and 21–300 nm), which constituted up to 100% of the total river P discharge depending on hydromorphology. For the small size fraction, variations in P concentrations correlated with Al variations; in addition, a high Fe presence in both fractions was accompanied by high P concentrations. Moreover, DOM was detected with P in the presence of Fe and Al, suggesting that Fe and Al are carriers of P and associated with organic matter. The developed methodology enables the inputs and source regions of fine colloidal and nanoparticulate fractions within a small river of a headwater catchment to be traced and conceptually defined for the first time.

Biogeochemical cycling of elements largely occurs in dissolved state, but many elements may also be bound to natural nanoparticles (NNP, 1–100 nm) and fine colloids (100–450 nm). We examined the hypothesis that the size and composition of stream water NNP and colloids vary systematically across Europe. To test this hypothesis, 96 stream water samples were simultaneously collected in 26 forested headwater catchments along two transects across Europe. Three size fractions (~1–20 nm, >20–60 nm, and >60 nm) of NNP and fine colloids were identified with Field Flow Fractionation coupled to inductively coupled plasma mass spectrometry and an organic carbon detector. The results showed that NNP and fine colloids constituted between 2 ± 5% (Si) and 53 ± 21% (Fe; mean ± SD) of total element concentrations, indicating a substantial contribution of particles to element transport in these European streams, especially for P and Fe. The particulate contents of Fe, Al, and organic C were correlated to their total element concentrations, but those of particulate Si, Mn, P, and Ca were not. The fine colloidal fractions >60 nm were dominated by clay minerals across all sites. The resulting element patterns of NNP <60 nm changed from North to South Europe from Fe‐ to Ca‐dominated particles, along with associated changes in acidity, forest type, and dominant lithology.

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