Fine root foraging strategies in Norway spruce forests across a European climate gradient

Fine root acclimation to different environmental conditions is crucial for growth and sustainability of forest trees. Relatively small changes in fine root standing biomass (FRB), morphology or mycorrhizal symbiosis may result in a large change in forest carbon, nutrient and water cycles. We elucidated the changes in fine root traits and associated ectomycorrhizal (EcM) fungi in 12 Norway spruce stands across a climatic and N deposition gradient from subarctic‐boreal to temperate regions in Europe (68°N–48°N). We analysed the standing FRB and the ectomycorrhizal root tip biomass (EcMB, g m^?2) simultaneously with measurements of the EcM root morphological traits (e.g. mean root length, root tissue density (RTD), N% in EcM roots) and frequency of dominating EcM fungi in different stands in relation to climate, soil and site characteristics. Latitude and N deposition explained the greatest proportion of variation in fine root traits. EcMB per stand basal area (BA) increased exponentially with latitude: by about 12.7 kg m^?2 with an increase of 10° latitude from southern Germany to Estonia and southern Finland and by about 44.7 kg m^?2 with next latitudinal 10° from southern to northern Finland. Boreal Norway spruce forests had 4.5 to 11 times more EcM root tips per stand BA, and the tips were 2.1 times longer, with 1.5 times higher RTD and about 1/3 lower N concentration. There was 19% higher proportion of root tips colonized by long‐distance exploration type forming EcM fungi in the southern forests indicating importance of EcM symbiont foraging strategy in fine root nutrient acquisition. In the boreal zone, we predict ca. 50% decrease in EcMB per stand BA with an increase of 2 °C annual mean temperature. Different fine root foraging strategies in boreal and temperate forests highlight the importance of complex studies on respective regulatory mechanisms in changing climate.     

Oxidation of atmospheric methane in Northern European soils, comparison with other ecosystems, and uncertainties in the global terrestrial sink

This paper reports the range and statistical distribution of oxidation rates of atmospheric CH₄ in soils found in Northern Europe in an international study, and compares them with published data for various other ecosystems. It reassesses the size, and the uncertainty in, the global terrestrial CH₄ sink, and examines the effect of land‐use change and other factors on the oxidation rate. Only soils with a very high water table were sources of CH₄; all others were sinks. Oxidation rates varied from 1 to nearly 200 μg CH₄ m^?2 h^?1; annual rates for sites measured for ≥1 y were 0.1–9.1 kg CH₄ ha^?1 y^?1, with a log‐normal distribution (log‐mean ≈ 1.6 kg CH₄ ha^?1 y^?1). Conversion of natural soils to agriculture reduced oxidation rates by two‐thirds –‐ closely similar to results reported for other regions. N inputs also decreased oxidation rates. Full recovery of rates after these disturbances takes > 100 y. Soil bulk density, water content and gas diffusivity had major impacts on oxidation rates. Trends were similar to those derived from other published work. Increasing acidity reduced oxidation, partially but not wholly explained by poor diffusion through litter layers which did not themselves contribute to the oxidation. The effect of temperature was small, attributed to substrate limitation and low atmospheric concentration. Analysis of all available data for CH₄ oxidation rates in situ showed similar log‐normal distributions to those obtained for our results, with generally little difference between different natural ecosystems, or between short‐and longer‐term studies. The overall global terrestrial sink was estimated at 29 Tg CH₄ y^?1, close to the current IPCC assessment, but with a much wider uncertainty range (7 to > 100 Tg CH₄ y^?1). Little or no information is available for many major ecosystems; these should receive high priority in future research.     

Nutritional influence on childhood development and genetic control of adolescent growth of Quechuas and Mestizos from the Peruvian Lowlands

The growth in height of 1,202 Quechua and Mestizo children aged 6 to 19 years of the province of Lamas in the Peruvian Eastern Lowlands was studied. As shown by evaluations of ABO, Rh systems, and skin reflectance measurements, the Quechuas are genetically different from the Mestizos. The heights of Quechuas and Mestizos were matched for nutritional status based on measurements of subcutaneous fat and body muscle. The study indicates that: (1) during childhood, Quechuas and Mestizos matched for the same nutritional status attain similar heights; (2) during adolescence (or after the age of 11 years), the Mestizos are significantly taller than the Quechuas of the same nutritional status; 3) during childhood, the relative difference in height between Quechuas and Mestizos matched for the same nutritional status is less than the difference between Quechuas (or Mestizos) of the same genetic composition characterized by good and poor nutritional status. These findings suggest that the influence of environmental factors, such as nutrition, have a greater influence in producing differences in body size during childhood than during adolescence. Conversely, the present findings support the hypothesis that the influence of genetic factors on body size are greater during adolescence than during childhood. However, comparison of adolescent samples of similar genetic composition (whether they be Quechuas or Mestizos), characterized by good and poor nutritional status, reveal large differences in height, suggesting that under conditions of malnutrition, the genetic control of growth is diminished.     

Does long-term soil warming affect microbial element limitation? A test by short-term assays of microbial growth responses to labile C,N and P additions

Increasing global temperatures have been reported to accelerate soil carbon (C) cycling, but also to promote nitrogen (N) and phosphorus (P) dynamics in terrestrial ecosystems. However, warming can differentially affect ecosystem C, N and P dynamics, potentially intensifying elemental imbalances between soil resources, plants and soil microorganisms. Here, we investigated the effect of long-term soil warming on microbial resource limitation, based on measurements of microbial growth (¹⁸O incorporation into DNA) and respiration after C, N and P amendments. Soil samples were taken from two soil depths (0–10, 10–20 cm) in control and warmed (>14 years warming, +4°C) plots in the Achenkirch soil warming experiment. Soils were amended with combinations of glucose-C, inorganic/organic N and inorganic/organic P in a full factorial design, followed by incubation at their respective mean field temperatures for 24 h. Soil microbes were generally C-limited, exhibiting 1.8-fold to 8.8-fold increases in microbial growth upon C addition. Warming consistently caused soil microorganisms to shift from being predominately C limited to become C-P co-limited. This P limitation possibly was due to increased abiotic P immobilization in warmed soils. Microbes further showed stronger growth stimulation under combined glucose and inorganic nutrient amendments compared to organic nutrient additions. This may be related to a prolonged lag phase in organic N (glucosamine) mineralization and utilization compared to glucose. Soil respiration strongly positively responded to all kinds of glucose-C amendments, while responses of microbial growth were less pronounced in many of these treatments. This highlights that respiration–though easy and cheap to measure—is not a good substitute of growth when assessing microbial element limitation. Overall, we demonstrate a significant shift in microbial element limitation in warmed soils, from C to C-P co-limitation, with strong repercussions on the linkage between soil C, N and P cycles under long-term warming.     

Responses of CO2 Exchange and Primary Production of the Ecosystem Components to Environmental Changes in a Mountain Peatland

The complexity of natural ecological systems presents challenges for predicting the impact of global environmental changes on ecosystem structure and function. Grouping of plants into functional types, that is, groups of species sharing traits that govern their mechanisms of response to environmental perturbations, reduce the complexity of species diversity to a few key plant types for better understanding of ecosystem responses. Chambers were used to measure CO₂ exchange in grass and moss growing together in a mountain peatland in southern Germany to assess variations in their response to environmental changes and how they influence ecosystem CO₂ exchange. Parameter fits and comparison for net ecosystem exchange (NEE) in two ecosystem components were conducted using an empirical hyperbolic light response model. Annual green biomass production was 320 and 210 g dwt m ⁻², whereas mean maximum NEE was –10.0 and –5.0 μmol m ⁻² s ⁻¹ for grass and moss, respectively. Grass exhibited higher light use efficiency (α) and maximum gross primary production [(β+γ)₂₀₀₀]. Leaf area index explained 93% of light use and 83% of overall production by the grass. Peat temperature at 10-cm depth explained more than 80% of the fluctuations in ecosystem respiration (R _eco). Compared to grass, moss NEE was more sensitive to ground water level (GWL) draw-down and hence could be more vulnerable to changes in precipitation that result in GWL decline and may be potentially replaced by grass and other vegetation that are less sensitive. Author’s Contribution  Werner Borken conceived the study. Ai Nishiwaki, Margerete Wartinger, G. Lischeid and Zaman Hussain conducted measurements. Jan Muhr helped with the methodologies and result discussion. Dennis O. Otieno designed and conducted measurements and wrote the paper.     

N2O emission in a Norway spruce forest due to soil frost: concentration and isotope profiles shed a new light on an old story

In mountain regions of Central Europe an increase of soil frost periods is predicted for this century due to reduced snow fall. To investigate the effects of freezing and thawing on soil N₂O fluxes in a mature Norway spruce forest in the mountainous Fichtelgebirge, Germany, the natural snow cover on three experimental plots was removed to induce soil frost. Three plots with natural snow cover served as controls. Soil N₂O fluxes were recorded in biweekly to monthly intervals during the frost and subsequent thawing period of the below-average cold winter in 2005/2006 and in the above-average warm winter in 2006/2007. In addition, N₂O concentrations and isotope signatures in soil air were measured along soil profiles in six different depths (from 6 to 70 cm). The soil of the snow removal plots was frozen down to 15 cm depth from January to April 2006 while the soil of control plots remained unfrozen under snow cover. Both soil freezing and thawing resulted in almost tenfold enhanced N₂O fluxes on snow removal plots contributing 84% to annual N₂O emissions. In the subsequent winter without soil frost no effects were observed. Vertical gradients of N₂O concentrations together with isotope abundance suggest that the subsoil of all plots was a probably weak, but continuous N₂O source throughout the year. Isotope signatures and N₂O concentration gradients in the soil profile indicate that microbial N₂O production and reduction of N₂O to N₂ did not or just marginally occur in frozen soil layers of the snow removal plots. Consequently, elevated N₂O fluxes in the late winter were attributed to the release of accumulated N₂O originating from the subsoil. At unfrozen soil, however, N₂O emissions were reduced due to a shift of the N₂O production-consumption ratio towards more consumption in the topsoil of both the control and snow removal plots. These findings contradict the general assumption that N₂O production in the organic layer is responsible for bursts of N₂O due to soil frost.     

Ammonium,nitrate and dissolved organic nitrogen in seepage water as affected by compost amendment to European beech,Norway spruce,and Scots pine forests

Fertilization of nutrient-depleted and degraded forest soils may be required to sustain utilization of forests. In some European countries, the application of composts may now be an alternative to the application of inorganic fertilizers because commercial compost production has increased and compost quality has been improved. There is, however, concern that compost amendments may cause increased leaching of nitrogen, trace metals and toxic organic compounds to groundwater. The objective of this study was to assess the risk of ammonium (NH₄ ⁺), nitrate (NO₃ ^–) and dissolved organic nitrogen (DON) leaching following a single compost application to silty and sandy soils in mature beech (Fagus sylvatica L.), pine (Pinus silvestris L.) and spruce (Picea abies Karst.) forests at Solling and Unterlüß in Lower Saxony, Germany. Mature compost from separately collected organic household waste was applied to the soil surface at a rate of 6.3 kg m^–2 in the summer of 1997 and changes in NH₄ ⁺, NO₃ ^– and DON concentrations in throughfall and soil water at 10 and 100 cm soil depths were determined for 32 months. The spruce forests had the highest N inputs by throughfall water and the highest N outputs in both the control and compost plots compared with the pine and beech forests. Overall, the differences in total N outputs at 100 cm soil depth between the control and compost plots ranged between 0.3 and 11.2 g N m^–2 for the entire 32-month period. The major leaching of these amounts occurred during the first 17 months after compost amendments, but there was no significant difference in total N outputs (–0.2 to 1.8 g N m^–2) between the control and compost plots during the remaining 15 months. Most of the mineral soils acted as a significant sink for NO₃ ^– and DON as shown by a reduction of their outputs from 10 to 100 cm depth. Based on these results, we conclude that application of mature compost with high inorganic N contents could diminish the groundwater quality in the first months after the amendments. A partial, moderate application of mature compost with low inorganic N content to nutrient depleted forest soils can minimize the risk of NO₃ ^– leaching.     

Dynamics of dissolved organic <Superscript>14</Superscript>C in throughfall and soil solution of a Norway spruce forest

Dissolved organic carbon (DOC) is an important component of the C cycle in forest ecosystems, but dynamics and origin of DOC in throughfall and soil solution are yet poorly understood. In a 2-year study, we analyzed the radiocarbon signature of DOC in throughfall and soil solution beneath the Oa horizon and at 90 cm depth in a Norway spruce forest on a Podzol soil. A two-pool mixing model revealed that throughfall DOC comprised mainly biogenic C, i.e. recently fixed C, from canopy leaching and possibly other sources. The contribution of fossil DOC from atmospheric deposition to throughfall DOC was on average 6% with maxima of 8–11% during the dormant season. In soil solution from the Oa horizon, DO¹⁴C signature was highly dynamic (range from −8‰ to +103‰), but not correlated with DOC concentration. Radiocarbon signatures suggest that DOC beneath the Oa horizon originated mainly from occluded and mineral associated organic matter fractions of the Oa horizon rather than from the Oi or Oe horizon. Relatively old C was released in the rewetting phase following a drought period in the late summer of 2006. In contrast, the DO¹⁴C signature indicated the release of younger C throughout the humid year 2007. In soil solutions from 90 cm depth, DO¹⁴C signatures were also highly dynamic (−127‰ to +3‰) despite constantly low DOC concentrations. Similar to the Oa horizon, the lowest DO¹⁴C signature at 90 cm depth was found after the rewetting phase in the late summer of 2006. Because of the variation in the DO¹⁴C signatures at this depth, we conclude that DOC was not equilibrated with the surrounding soil, but also originated from overlaying soil horizons. The dynamics of DO¹⁴C in throughfall and soil solution suggest that the sources of DOC are highly variable in time. Extended drought periods likely have a strong influence on release and translocation of DOC from relatively old and possibly stabilized soil organic matter fractions. Temporal variations as well as the input of fossil DOC needs to be considered when calibrating DOC models based on DO¹⁴C signatures.     

Carbon fluxes in coniferous and deciduous forest soils

Aims

Our aims were to identify responsible factors for the site-to-site variability in soil CO₂ efflux and to assess the sources of soil CO₂ of different forest types on a regional scale.

Methods

Soil CO₂ effluxes were measured over 1–4 years in four coniferous and three deciduous forests of Bavaria, Germany, and related to climate, soil properties and forest productivity. Total belowground carbon allocation (TBCA) was assessed using soil CO₂ effluxes and aboveground litterfall. Additionally, CO₂ production of organic layers was examined over 10 months under constant conditions in an incubation experiment.

Results

Annual soil CO₂ effluxes were not different among the forest sites, but predicted effluxes at a given temperature of 10°C revealed some significant differences and correlated with the phosphorous stock of the organic layers. The incubation study indicated 50% faster decomposition of organic layers from deciduous than from coniferous forests. TBCA related to soil CO₂ efflux was smaller in the deciduous than in the coniferous forests. The ratio of TBCA to soil CO₂ efflux was positively correlated with the C stock of organic layers.

Conclusions

Our results suggest that marked differences in site characteristics have little impact on soil CO₂ effluxes at the regional scale, but the contribution of soil CO₂ sources varies among the forest types.