Biological nitrogen fixation in logging residue piles of different tree species after final felling.

Logging residues influence the nitrogen cycling processes that play a key role in risks for nitrogen losses from the ecosystem after the clear cut. Therefore, our aim was to identify the potential ability of logging residues to gain external nitrogen via biological nitrogen fixation. We measured biological nitrogen fixation as nitrogenase activity in logging residues of three different tree species (Norway spruce (Picea abies (L.) Karst.), Scots pine (Pinus sylvestris L.), and silver birch (Betula pendula Roth.). The study site was located in south-eastern Finland and was clear cut in 2014 and piles of logging residues were established. Sampling was performed in June 2016, September 2018, and August 2019 and nitrogenase activity in branches and needles or leaves was measured using the acetylene (C2H2) reduction assay. Nitrogenase activity (ethylene production) was shown in all residue types. Nitrogenase activity tended to be higher in branches than in needles or leaves and in coniferous residues than in birch. C-to-N ratios were higher in branches than in needles/leaves and in coniferous residues than in birch. Our results indicate that logging residues can acquire external nitrogen from the atmosphere via biological nitrogen fixation and can thus bring nitrogen to the forest ecosystem and substitute some part of the N losses occurring when residues are retained at the site after clear cutting.

Logging residue piles of Norway spruce, Scots pine and silver birch in a clear-cut: Effects on nitrous oxide emissions and soil percolate water nitrogen.

We analysed how logging residue (LR) piles of common tree species in Finland, Norway spruce (Picea abies (L.) H. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth), affect nitrogen (N) losses in forest soil after final felling. A Norway spruce dominated stand was clear-cut and followed by two experimental setups to study the nitrous oxide (N2O) emissions and leaching of carbon (C) and N. Experiments consisted of four treatments: tree species treatments consisting of 40 kg m-2 of LR and a control treatment without residues. The C losses were monitored as dissolved organic carbon (DOC), the N losses as ammonium (NH4-N), nitrate (NO3-N) and dissolved organic nitrogen (DON) fluxes and concentrations in soil percolation waters and the N2O emissions as fluxes from the forest soil to the atmosphere. In addition the soil temperatures, the molecular size distribution of the DOC from the soil percolation waters and the origin of the N2O production were determined. The LR piles lowered the soil temperatures and, especially those of birch, increased the concentrations of NO3-N in the soil percolation waters already 1 year after the establishment of the piles. The LR piles increased the NH4-N concentrations. The smallest molecular size fraction (<1 kD) of DOC predominated in all treatments. The N2O fluxes peaked under the piles during the second and third growing seasons; however, the inconsistent fluxes tended to be low. The production of N2O was driven by both nitrification and denitrification processes, the proportion depending on the tree species. Our results indicate that LR piles accelerate N losses 1 year after the clear-cutting, especially NO3-N, which predominates in the soil percolation waters under the birch residues, whereas spruce residues tend to stimulate N2O emissions longer. These results have implications for sustainable forest management practices and nutrition of regrowing vegetation.

Artificial recharge of groundwater through sprinkling infiltration: impacts on forest soil and the nutrient status and growth of Scots pine.

We studied the chemical changes in forest soil and the effects on Scots pine trees caused by continuous sprinkling infiltration over a period of two years, followed by a recovery period of two years. Infiltration increased the water input onto the forest soil by a factor of approximately 1000. After one year of infiltration, the pH of the organic layer had risen from about 4.0 to 6.7. The NH(4)-N concentration in the organic layer increased, most probably due to the NH(4) ions in the infiltration water, as the net N mineralization rate did not increase. Sprinkling infiltration initiated nitrification in the mineral soil. Macronutrient concentrations generally increased in the organic layer and mineral soil. An exception, however, was the concentration of extractable phosphorus, which decreased strongly during the infiltration period and did not show a recovery within two years. The NO(3)-N and K concentrations had reverted back to their initial level during the two-year recovery period, while the concentrations of Ca, Mg and NH(4)-N were still elevated. Nutrient concentrations in the pine needles increased on the infiltrated plots. However, the needle P concentration increased, despite the decrease in plant-available P in the soil. Despite the increase in the nutrient status, there were some visible signs of chlorosis in the current-year needles after two years of infiltration. The radial growth of the pines more than doubled on the infiltrated plots, which suggests that the very large increase in the water input onto the forest floor had no adverse effect on the functioning of the trees. However, a monitoring period of four years is not sufficient for detecting potential long term detrimental effects on forest trees.

N2 fixation associated with the bryophyte layer is suppressed by low levels of nitrogen deposition in boreal forests.

Biological fixation of atmospheric nitrogen (N2) by bryophyte-associated cyanobacteria is an important source of plant-available N in the boreal biome. Information on the factors that drive biological N2 fixation (BNF) rates is needed in order to understand the N dynamics of forests under a changing climate. We assessed the potential of several cryptogam species (the feather mosses Hylocomium splendens and Pleurozium schreberi, a group of Dicranum bryophytes, two liverworts, and Cladina lichens) to serve as associates of cyanobacteria or other N2-fixing bacteria (diazotrophs) using acetylene reduction assay (ARA). We tested the hypotheses that the legacy of chronic atmospheric N deposition reduces BNF in the three bryophyte species, sampled from 12 coniferous forests located at latitudes 60-68° N in Finland. In addition, we tested the effect of moisture and temperature on BNF. All species studied showed a BNF signal in the north, with the highest rates in feather mosses. In moss samples taken along the north-south gradient with an increasing N bulk deposition from 0.8 to 4.4 kg ha-1 year-1, we found a clear decrease in BNF in both feather mosses and Dicranum group. BNF turned off at N deposition of 3-4 kg ha-1 year-1. Inorganic N (NH4-N + NO3-N) best predicted the BNF rate among regression models with different forms of N deposition as explanatory variables. However, in southern spruce stands, tree canopies modified the N in throughfall so that dissolved organic N (DON) leached from canopies compensated for inorganic N retained therein. Here, both DON and inorganic N negatively affected BNF in H. splendens. In laboratory experiments, BNF increased with increasing temperature and moisture. Our results suggest that even relatively low N deposition suppresses BNF in bryophyte-associated diazotrophs. Further, BNF could increase in northern low-deposition areas, especially if climate warming leads to moister conditions, as predicted.

Tannins and Their Complex Interaction with Different Organic Nitrogen Compounds and Enzymes: Old Paradigms versus Recent Advances.

Tannins, an abundant group of plant secondary compounds, raise interest in different fields of science, owing to their unique chemical characteristics. In chemical ecology, tannins play a crucial role in plant defense against pathogens, herbivores, and changing environmental conditions. In the food industry and in medicine, tannins are important because of their proven positive effect on human health and disease treatment. Such wide interests fueled studies on tannin chemistry, especially on their flagship ability to precipitate proteins. In this Review, we expand the basic knowledge on tannin chemistry to the newest insights from the field. We focus especially on tannin reactions with different non-protein organic N compounds, as well as the complex interactions of tannins with enzymes, resulting in either an increase or decrease in enzyme activity.

Degradability of dissolved soil organic carbon and nitrogen in relation to tree species.

The degradability and chemical characteristics of water-extractable dissolved organic carbon (DOC) and nitrogen (DON) from the humus layer of silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) stands were compared in short-term incubation of soil solutions. For all extracts the degradation of DOC and DON was low (12-17% loss) and increased in the order: birch, spruce and pine. In the humus layer under pine a relatively larger pool of rapidly degrading dissolved soil organic matter (DOM) was indicated by the [3H]thymidine incorporation technique, which measures the availability of DOM to bacteria. The degradation of DOC was explained by a decrease in the hydrophilic fraction. For DON, however, both the hydrophilic and hydrophobic fractions tended to decrease during incubation. No major differences in concentrations of hydrophilic and hydrophobic fractions were detected between tree species. Molecular size distribution of DOC and DON, however, revealed slight initial differences between birch and conifers as well as a change in birch extract during incubation. The depletion of very rapidly degrading fractions (e.g., root exudates and compounds from the litter) may explain the low degradability of DOM in the humus layer under birch.

Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests.

It has been recognized for a long time that the overstorey composition of a forest partly determines its biological and physical-chemical functioning. Here, we review evidence of the influence of evergreen gymnosperm (EG) tree species and deciduous angiosperm (DA) tree species on the water balance, physical-chemical soil properties and biogeochemical cycling of carbon and nutrients. We used scientific publications based on experimental designs where all species grew on the same parent material and initial soil, and were similar in stage of stand development, former land use and current management. We present the current state of the art, define knowledge gaps, and briefly discuss how selection of tree species can be used to mitigate pollution or enhance accumulation of stable organic carbon in the soil. The presence of EGs generally induces a lower rate of precipitation input into the soil than DAs, resulting in drier soil conditions and lower water discharge. Soil temperature is generally not different, or slightly lower, under an EG canopy compared to a DA canopy. Chemical properties, such as soil pH, can also be significantly modified by taxonomic groups of tree species. Biomass production is usually similar or lower in DA stands than in stands of EGs. Aboveground production of dead organic matter appears to be of the same order of magnitude between tree species groups growing on the same site. Some DAs induce more rapid decomposition of litter than EGs because of the chemical properties of their tissues, higher soil moisture and favourable conditions for earthworms. Forest floors consequently tend to be thicker in EG forests compared to DA forests. Many factors, such as litter lignin content, influence litter decomposition and it is difficult to identify specific litter-quality parameters that distinguish litter decomposition rates of EGs from DAs. Although it has been suggested that DAs can result in higher accumulation of soil carbon stocks, evidence from field studies does not show any obvious trend. Further research is required to clarify if accumulation of carbon in soils (i.e. forest floor + mineral soil) is different between the two types of trees. Production of belowground dead organic matter appears to be of similar magnitude in DA and EG forests, and root decomposition rate lower under EGs than DAs. However there are some discrepancies and still are insufficient data about belowground pools and processes that require further research. Relatively larger amounts of nutrients enter the soil-plant biogeochemical cycle under the influence of EGs than DAs, but recycling of nutrients appears to be slightly enhanced by DAs. Understanding the mechanisms underlying forest ecosystem functioning is essential to predicting the consequences of the expected tree species migration under global change. This knowledge can also be used as a mitigation tool regarding carbon sequestration or management of surface waters because the type of tree species affects forest growth, carbon, water and nutrient cycling.

The community of needle endophytes reflects the current physiological state of Norway spruce.

This study investigated fungal endophytes in the needles of Norway spruce (Picea abies) cuttings in relation to host tree growth. We also determined the prevalence of endophytes in needles incubated for six months. The cuttings originated from clonal origins showing slow- and fast-growth in long-term field trials but the heritable differences in growth rate were not yet detected among the studied cutting. Endophytes were isolated from surface-sterilized needles with culture-free DNA techniques. No significant differences were observed between endophyte communities of slow- and fast-growing clonal origins. However, the endophyte community correlated with the current growth rate of cuttings suggesting that endophytes reflect short- rather than long-term performance of a host. The concentration of condensed tannins was similar in slow- and fast-growing clonal origins but it showed a negative relationship with endophyte species richness, implying that these secondary compounds may play an important role in spruce tolerance against fungal infections. More than a third of endophyte species were detected in both fresh and decomposing needles, indicating that many needle endophytes are facultative saprotrophs. Several potentially pathogenic fungal species were also found within the community of saprotrophic endophytes.

Proteins as nitrogen source for plants: a short story about exudation of proteases by plant roots.

Interest in the problem of plant nitrogen nutrition is increasing. Certain plants can use not only inorganic nitrogen, but also intact amino acids and short peptides. According to our studies, the roots of several agricultural and wild-living plants are able to exude proteases and by using them to create a pool of accessible N. This mini-review offers an overview of the problem of protease exudation by plant roots and its potential role in plant nitrogen nutrition.

Degradation of proteins by enzymes exuded by Allium porrum roots - a potentially important strategy for acquiring organic nitrogen by plants.

Nitrogen is one of the crucial elements that regulate plant growth and development. It is well-established that plants can acquire nitrogen from soil in the form of low-molecular-mass compounds, namely nitrate and ammonium, but also as amino acids. Nevertheless, nitrogen in the soil occurs mainly as proteins or proteins complexed with other organic compounds. Proteins are believed not to be available to plants. However, there is increasing evidence to suggest that plants can actively participate in proteolysis by exudation of proteases by roots and can obtain nitrogen from digested proteins. To gain insight into the process of organic nitrogen acquisition from proteins by leek roots (Allium porrum L. cv. Bartek), casein, bovine serum albumin and oxidized B-chain of insulin were used; their degradation products, after exposure to plant culture medium, were studied using liquid chromatography-mass spectrometry (LC-MS). Casein was degraded to a great extent, but the level of degradation of bovine serum albumin and the B-chain of insulin was lower. Proteases exuded by roots cleaved proteins, releasing low-molecular-mass peptides that can be taken up by roots. Various peptide fragments produced by digestion of the oxidized B-chain of insulin suggested that endopeptidase, but also exopeptidase activity was present. After identification, proteases were similar to cysteine protease from Arabidopsis thaliana. In conclusion, proteases exuded by roots may have great potential in the plant nitrogen nutrition.