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Forests play a crucial role in global carbon cycling by absorbing and storing significant amounts of atmospheric carbon dioxide. Although boreal forests contribute to approximately 45% of the total forest carbon sink, tree growth and soil carbon sequestration are constrained by nutrient availability. Here, we examine if long-term nutrient input enhances tree productivity and whether this leads to carbon storage or whether stimulated microbial decomposition of organic matter limits soil carbon accumulation. Over six decades, nitrogen, phosphorus, and calcium were supplied to a Pinus sylvestris-dominated boreal forest. We found that nitrogen fertilization alone or together with calcium and/or phosphorus increased tree biomass production by 50% and soil carbon sequestration by 65% compared to unfertilized plots. However, the nonlinear relationship observed between tree productivity and soil carbon stock across treatments suggests microbial regulation. When phosphorus was co-applied with nitrogen, it acidified the soil, increased fungal biomass, altered microbial community composition, and enhanced biopolymer degradation capabilities. While no evidence of competition between ectomycorrhizal and saprotrophic fungi has been observed, key functional groups with the potential to reduce carbon stocks were identified. In contrast, when nitrogen was added without phosphorus, it increased soil carbon sequestration because microbial activity was likely limited by phosphorus availability. In conclusion, the addition of nitrogen to boreal forests may contribute to global warming mitigation, but this effect is context dependent., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Tree growth in boreal forests is driven by ectomycorrhizal fungal mobilisation of organic nitrogen and mineral nutrients in soils with discrete organic and mineral horizons. However, there are no studies of how ectomycorrhizal mineral weathering and organic nitrogen mobilisation processes are integrated across the soil profile. We studied effects of organic matter (OM) availability on ectomycorrhizal functioning by altering the proportions of natural organic and mineral soil in reconstructed podzol profiles containing Pinus sylvestris plants, using 13 CO 2 pulse labelling, patterns of naturally occurring stable isotopes ( 26 Mg and 15 N) and high-throughput DNA sequencing of fungal amplicons. Reduction in OM resulted in nitrogen limitation of plant growth and decreased allocation of photosynthetically derived carbon and mycelial growth in mineral horizons. Fractionation patterns of 26 Mg indicated that magnesium mobilisation and uptake occurred primarily in the deeper mineral horizon and was driven by carbon allocation to ectomycorrhizal mycelium. In this horizon, relative abundance of ectomycorrhizal fungi, carbon allocation and base cation mobilisation all increased with increased OM availability. Allocation of carbon through ectomycorrhizal fungi integrates organic nitrogen mobilisation and mineral weathering across soil horizons, improving the efficiency of plant nutrient acquisition. Our findings have fundamental implications for sustainable forest management and belowground carbon sequestration., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

A key weakness in current Earth System Models is the representation of thermal acclimation of photosynthesis in response to changes in growth temperatures. Previous studies in boreal and temperate ecosystems have shown leaf-scale photosynthetic capacity parameters, the maximum rates of carboxylation (V cmax ) and electron transport (J max ), to be positively correlated with foliar nitrogen (N) content at a given reference temperature. It is also known that V cmax and J max exhibit temperature optima that are affected by various environmental factors and, further, that N partitioning among the foliar photosynthetic pools is affected by N availability. However, despite the strong recent anthropogenic influence on atmospheric temperatures and N deposition to forests, little is known about the role of foliar N contents in controlling the photosynthetic temperature responses. In this study, we investigated the temperature dependencies of V cmax and J max in 1-year-old needles of mature boreal Pinus sylvestris (Scots pine) trees growing under low and high N availabilities in northern Sweden. We found that needle N status did not significantly affect the temperature responses of V cmax or J max when the responses were fitted to a peaked function. If such N insensitivity is a common tree trait it will simplify the interpretation of the results from gradient and multi-species studies, which commonly use sites with differing N availabilities, on temperature acclimation of photosynthetic capacity. Moreover, it will simplify modeling efforts aimed at understanding future carbon uptake by precluding the need to adjust the shape of the temperature response curves to variation in N availability., (© 2017 Scandinavian Plant Physiology Society.)

We report results from long-term simulated external nitrogen (N) input experiments in three northern Pinus sylvestris forests, two of moderately high and one of moderately low productivity, assessing effects on annual net primary production (NPP) of woody mass and its interannual variation in response to variability in weather conditions. A sigmoidal response of wood NPP to external N inputs was observed in the both higher and lower productivity stands, reaching a maximum of ~65% enhancement regardless of the native site productivity, saturating at an external N input of 4-5 g N·m -2 ·yr -1 . The rate of increase in wood NPP and the N response efficiency (RE N , increase in wood NPP per external N input) were maximized at an external N input of ~3 g N·m -2 ·yr -1 , regardless of site productivity. The maximum RE N was greater in the higher productivity than the lower productivity stand (~20 vs. ~14 g C/g N). The N-induced enhancement of wood NPP and its RE N were, however, markedly contingent on climatic variables. In both of the higher and lower productivity stands, wood NPP increased with growing season precipitation (P), but only up to ~400 mm. The sensitivity of the response to P increased with increasing external N inputs. Increasing growing season temperature (T) somewhat increased the N-induced drought effect, whereas decreasing T reduced the drought effect. These responses of wood NPP infused a large temporal variation to RE N , making the use of a fixed value unadvisable. Based on these results, we suggest that regional climate conditions and future climate scenarios should be considered when modeling carbon sequestration in response to N deposition in boreal P. sylvestris, and possibly other forests., (© 2017 by the Ecological Society of America.)

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 δ(13)C and δ(18)O in host tree rings. After mistletoe removal, δ(13)C did not change in newly produced tree rings compared with tree rings in control trees (still infected with mistletoe), but δ(18)O 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 δ(13)C 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., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Ectomycorrhizal (ECM) symbiosis has been proposed to link plant photosynthesis and soil organic matter (SOM) decomposition through the production of fungal enzymes which promote SOM degradation and nitrogen (N) uptake. However, laboratory and field evidence for the existence of these processes are rare. Piloderma sp., a common ECM genus in boreal forest soil, was chosen as model mycorrhiza for this study. The abundance of Piloderma sp. was studied in root tips and soil over one growing season and in winter. Protease production was measured from ectomycorrhiza and soil solution in the field and pure fungal cultures. We also tested the effect of Piloderma olivaceum on host plant organic N nutrition in the laboratory. The results showed that Piloderma sp. was highly abundant in the field and produced extracellular proteases, which correlated positively with the gross primary production, temperature and soil respiration. In the laboratory, Piloderma olivaceum could improve the ability of Pinus sylvestris L. to utilize N from extragenous proteins. We suggest that ECM fungi, although potentially retaining N in their hyphae, are important in forest C and N cycling due to their ability to access proteinaeous N. As Piloderma sp. abundance appeared to be seasonally highly variable, recycling of fungal-bound N after hyphal death may therefore be of primary importance for the N cycling in boreal ecosystems.

Insights into how the simultaneous presence of organic and inorganic nitrogen (N) forms influences root absorption will help elucidate the relative importance of these N forms for plant nutrition in the field as well as for nursery cultivation of seedlings. Uptake of the individual N forms arginine, ammonium (NH4(+)) and nitrate (NO3(-)) was studied in Scots pine (Pinus sylvestris (L.)) seedlings supplied as single N sources and additionally in mixtures of NO3(-) and NH4(+) or NO3(-) and arginine. Scots pine seedlings displayed a strong preference for NH4(+)-N and arginine-N as compared with NO3(-)-N. Thus, NO3(-) uptake was generally low and decreased in the presence of NH4(+) in the high-concentration range (500 µM N), but not in the presence of arginine. Moreover, uptake of NO3(-) and NH4(+) was lower in seedlings displaying a high internal N status as a result of high N pre-treatment, while arginine uptake was high in seedlings with a high internal N status when previously exposed to organic N. These findings may have practical implications for commercial cultivation of conifers.

A main objective of restoration and afforestation at post-mining sites is establishing a long-term sustainable ecosystem which depends on adaptations of tree species and which in turn depends on the soil nutrient flux. The nutrient concentration (nitrogen (N), P, K, Ca, Mg, and sulfur (S)) of Scots pine needles was investigated in reclaimed mine soils (RMS) located at the following post-mining sites: a sand mine pit, spoil heap from a lignite mine, spoil heap from a S mine, and a carbonaceous spoil heap from an underground coal mine. The control plots were arranged on natural forest sites adjacent to the post-mining sites. A higher level of foliar nutrients was noted in the carbonaceous RMS, while lower levels were found in RMS on the spoil heap following lignite mining. The characteristics of the substrate were found to exert greater effect than mineral fertilization (performed at the onset of reclamation) on the tree stand characteristics, needle length and foliar nutrient concentration. While the soils and trees were most deficient in N, negative symptoms have not been noted to this date in tree stands at reclaimed mine sites. Trophic ratings were recommended based on statistical correlations and groupings between N and P contents in needles and needles length (mean length of 300 needles) while nutrient ratings were recommended from statistical differences and groupings of the RMS substrates.

In boreal forests, seedling establishment is limited by various factors including soil nitrogen (N) availability. Seedlings may absorb N from soil in a variety of inorganic and organic forms; however, the energy and thus carbohydrate requirements for uptake and assimilation of N vary with N source. We studied the importance of current photoassimilates for the acquisition and allocation of different N sources by Scots pine (Pinus sylvestris (L.)) seedlings. Girdling was used as a tool to impair phloem transport of photoassimilates, and hence gradually deprive roots of carbohydrates. Seedlings were cultivated in a greenhouse on equimolar N concentrations of one of the N sources-arginine, ammonium or nitrate-and then girdled prior to a pulse-chase uptake experiment with isotopically labeled N sources. Girdling proved to be efficient in decreasing levels of soluble sugars and starch in the roots. Uptake rate of arginine N was highest, intermediate for ammonium N and lowest for nitrate N. Moreover, the uptake of arginine N was unaffected by girdling, while the uptake of the two inorganic N sources decreased to 45-56% of the ungirdled controls. In arginine-treated seedlings, 95-96% of the acquired arginine N resided in the roots, whereas a significant shift in the N distribution toward the shoot was evident in girdled seedlings treated with inorganic N. This spatial shift was especially pronounced in nitrate-treated seedlings suggesting that the reduction and following incorporation into roots was limited by the availability of current photoassimilates. These results suggest that there are energetic benefits for seedlings to utilize organic N sources, particularly under circumstances where carbohydrate supply is limited. Hence, these putative benefits might be of importance for the survival and growth of seedlings when carbohydrate reserves are depleted in early growing season, or in light-limited environments, such as those sustained by continuous cover forestry systems.

We investigated the interaction of rhizospheric nitric oxide (NO) concentration (i.e. low, ambient or high) and soil nitrogen (N) availability (i.e. low or high) with organic and inorganic N uptake by fine roots of Pinus sylvestris L. seedlings by (15) N feeding experiments under controlled conditions. N metabolites in fine roots were analysed to link N uptake to N nutrition. NO affected N uptake depending on N source and soil N availability. The suppression of nitrate uptake in the presence of ammonium and glutamine was overruled by high NO. The effects of NO on N uptake with increasing N availability showed different patterns: (1) increasing N uptake regardless of NO concentration (i.e. ammonium); (2) increasing N uptake only with high NO concentration (i.e. nitrate and arginine); and (3) decreasing N uptake (i.e. glutamine). At low N availability and high NO nitrate accumulated in the roots indicating insufficient substrates for nitrate reduction or its storage in root vacuoles. Individual amino acid concentrations were negatively affected with increasing NO (i.e. asparagine and glutamine with low N availability, serine and proline with high N availability). In conclusion, this study provides first evidence that NO affects N uptake and metabolism in a conifer., (© 2012 Blackwell Publishing Ltd.)

We formulate a dynamic evolutionary optimization problem to predict the optimal pattern by which carbon (C) and nitrogen (N) are co-allocated to fine-root, leaf, and wood production, with the objective of maximizing height growth rate, year by year, in an even-aged stand. Height growth is maximized with respect to two adaptive traits, leaf N concentration and the ratio of fine-root mass to sapwood cross-sectional area. Constraints on the optimization include pipe-model structure, the C cost of N acquisition, and agreement between the C and N balances. The latter is determined by two models of height growth rate, one derived from the C balance and the other from the N balance; agreement is defined by identical growth rates. Predicted time-courses of maximized height growth rate accord with general observations. Across an N gradient, higher N availability leads to greater N utilization and net primary productivity, larger trees, and greater stocks of leaf and live wood biomass, with declining gains as a result of saturation effects at high N availability. Fine-root biomass is greatest at intermediate N availability. Predicted leaf and fine-root stocks agree with data from coniferous stands across Finland. Optimal C-allocation patterns agree with published observations and model analyses., (No claim to original US government works. New Phytologist © 2012 New Phytologist Trust.)

Global environmental change has serious implications for functional biodiversity in temperate and boreal forests. Trees depend on mycorrhizal fungi for nutrient uptake, but predicted increases in nitrogen availability may alter fungal communities. To address a knowledge gap regarding the effects of nitrogen availability on mycorrhizal communities at large scales, we examine the relationship between nitrogen and ectomycorrhizas in part of a European biomonitoring network of pine forest plots. Our analyses show that increased nitrogen reduces fungal diversity and causes shifts in mycorrhizal community composition across plots, but we do not find strong evidence that within-plot differences in nitrogen availability affect ectomycorrhizal communities. We also carry out exploratory analyses to determine the relative importance of other environmental variables in structuring mycorrhizal communities, and discuss the potential use of indicator species to predict nitrogen-induced shifts in fungal communities.

Variations in ectomycorrhizal (EcM) short root tips of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in 16 stands throughout Finland were studied, and their relationships with latitude, organic layer C:N ratio, temperature sum and foliage biomass were determined. There were no significant differences in EcM root tip frequency (number per milligram of fine roots) or root tip mass between tree species or between northern and southern sites. The EcM root tip number per unit area of the organic layer plus the 0-30 cm mineral soil layer varied between 0.8 and 2.4 million per m(2) for Norway spruce and between 0.7 and 2.9 million per m(2) for Scots pine, and it was higher in the northern Scots pine stands than in the southern Scots pine stands. Over 80% of the EcM root tips of both species were in the organic layer and the upper 0-10 cm mineral soil layer. We related EcM root tips to foliage mass because these two components are the most important functional units in boreal tree physiology. Both species, especially the Scots pine trees, had more EcM root tips in relation to foliage mass in northern Finland than in southern Finland. Scots pine trees had more EcM root tips in relation to foliage mass than Norway spruce in the same climatic region. The EcM root tip:foliage biomass ratio of Norway spruce was positively related to the C:N ratio in the organic layer, whereas that of Scots pine was negatively related to the temperature sum. The number of EcM root tips per milligram of fine root biomass was constant, implying that trees of both species increase nutrient uptake by increasing fine root production and hence their total number of EcM tips and the area of soil occupied by mycelia. Both tree species responded to nitrogen (N) deficiency by maintaining more EcM tips per foliage unit, and this may be related to a higher proportion of N uptake in an organic form.

Phenolic compounds often accumulate in foliar tissues of deciduous woody plants in response to previous insect defoliation, but similar responses have been observed infrequently in evergreen conifers. We studied the effects of defoliation on the foliar chemistry of Scots pine (Pinus sylvestris L.) and cocoon mass, and survival of the pine sawfly (Diprion pini L.). In two successive years, needles were excised early in the season leaving only the current-year shoot intact (defoliated trees); untreated entire shoots served as controls (control trees). A year after the second defoliation, pine sawfly larvae were transferred to the trees. Delayed induced resistance in Scots pine in response to defoliation was indicated by (1) reduced cocoon mass in defoliated trees and (2) increased concentrations of phenolics and soluble condensed tannins in the foliage of defoliated trees compared with controls. Myricetin-3-galactoside, which showed the strongest induced response (104% and 71% increase in current-year (C) and previous-year (C+1) needles) of the compounds analyzed, also entered the regression model explaining variation in sawfly performance. Other compounds that entered the model, e.g., (+)-catechin, showed weaker responses to defoliation than myricetin-3-galactoside. Hyperin, condensed tannins and quercitrin showed strong induced responses in C or C+1 needles, or both, but these compounds did not explain the variation in sawfly performance. Accumulation of phenolics is sometimes associated with the reduced foliage nitrogen (N) concentrations in deciduous trees, and our results suggest that this may also be the case in evergreen conifers. Based on the earlier findings that defoliation reduces needle N concentration and N deficiency results in the accumulation of the same phenolic compounds, i.e., myricetin and quercetin glycosides, and soluble condensed tannins, we suggest that the accumulation of phenolics in defoliated trees occurred in response to the reduced foliar N concentration.

Needles of 20-year-old Scots pine (Pinus sylvestris L.) saplings were studied in an ultraviolet (UV) exclusion field experiment (from 2000 to 2002) in northern Finland (67 degrees N). The chambers held filters that excluded both UV-B and UV-A, excluded UV-B only, transmitted all UV (control), or lacked filters (ambient). UV-B/UV-A exclusion decreased nitrate reductase (NR) activity of 1-year-old needles of Scots pines compared to the controls. The proportion of free amino acids varied in the range 1.08-1.94% of total proteins, and was significantly higher in needles of saplings grown under UV-B/UV-A exclusion compared to the controls or UV-B exclusion. NR activity correlated with air temperature, indicating a "chamber effect". The study showed that both UV irradiance and increasing temperature are significant modulators of nitrogen (N) metabolism in Scots pine needles.

Nitrogen (N) is essential for plant production, but N uptake imposes carbon (C) costs through maintenance respiration and fine-root construction, suggesting that an optimal C:N balance can be found. Previous studies have elaborated this optimum under exponential growth; work on closed canopies has focused on foliage only. Here, the optimal co-allocation of C and N to foliage, fine roots and live wood is examined in a closed forest stand. Optimal co-allocation maximizes net primary productivity (NPP) as constrained by stand-level C and N balances and the pipe model. Photosynthesis and maintenance respiration increase with foliar nitrogen concentration ([N]), and stand-level photosynthesis and N uptake saturate at high foliage and fine-root density. Optimal NPP increases almost linearly from low to moderate N availability, saturating at high N. Where N availability is very low or very high, the system resembles a functional balance with a steady foliage [N]; in between, [N] increases with N availability. Carbon allocation to fine roots decreases, allocation to wood increases, and allocation to foliage remains stable with increasing N availability. The predicted relationships between biomass density and foliage [N] are in reasonable agreement with data from coniferous stands across Finland. All predictions agree with our qualitative understanding of N effects on growth.

Our understanding of how saprotrophic and mycorrhizal fungi interact to re-circulate carbon and nutrients from plant litter and soil organic matter is limited by poor understanding of their spatiotemporal dynamics. In order to investigate how different functional groups of fungi contribute to carbon and nitrogen cycling at different stages of decomposition, we studied changes in fungal community composition along vertical profiles through a Pinus sylvestris forest soil. We combined molecular identification methods with 14C dating of the organic matter, analyses of carbon:nitrogen (C:N) ratios and 15N natural abundance measurements. Saprotrophic fungi were primarily confined to relatively recently (< 4 yr) shed litter components on the surface of the forest floor, where organic carbon was mineralized while nitrogen was retained. Mycorrhizal fungi dominated in the underlying, more decomposed litter and humus, where they apparently mobilized N and made it available to their host plants. Our observations show that the degrading and nutrient-mobilizing components of the fungal community are spatially separated. This has important implications for biogeochemical studies of boreal forest ecosystems.

Although an increasing number of studies show that many plant species have the capacity to take up amino acids from exogenous sources, the importance of such uptake for plant nitrogen nutrition is largely unknown. Moreover, little is known regarding metabolism and distribution of amino acid-N following uptake or of the regulation of these processes in response to plant nitrogen status. Here results are presented from a study following uptake, metabolism, and distribution of nitrogen from NO(3)(-) NH(4)(+), Glu, or Ala in Scots pine (Pinus sylvestris L). In a parallel experiment, Ala uptake, processing, and shoot allocation were also monitored following a range of pretreatments intended to alter plant C- and N-status. Uptake data, metabolite profiles, N fluxes through metabolite pools and tissues, as well as alanine aminotransferase activity are presented. The results show that uptake of the organic N sources was equal to or larger than NH(4)(+) uptake, while NO(3)(-) uptake was comparatively low. Down-regulation of Ala uptake in response to pretreatments with NH(4)NO(3) or methionine sulphoximine (MSX) indicates similarities between amino acid and inorganic N uptake regulation. N derived from amino acid uptake exhibited a rapid flux through the amino acid pool following uptake. Relative shoot allocation of amino acid-N was equal to that of NH(4)(+) but smaller than for NO(3)(-) Increased N status as well as MSX treatment significantly increased relative shoot allocation of Ala-N suggesting that NH(4)(+) may have a role in the regulation of shoot allocation of amino acid-N.