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.     

Long-term effects on the nitrogen budget of a short-rotation grey alder (Alnus incana (L.) Moench) forest on abandoned agricultural land

Short-rotation energy forestry is one of the potential ways for management of abandoned agricultural areas. It helps sequestrate carbon and mitigate human-induced climate changes. Owing to symbiotic dinitrogen (N₂) fixation by actinomycetes and the soil fertilizing capacity and fast biomass growth of grey alders, the latter can be suitable species for short-rotation forestry. In our study of a young grey alder stand (Alnus incana (L.) Moench) on abandoned arable land in Estonia we tested the following hypotheses: (1) afforestation of abandoned agricultural land by grey alder significantly affects the soil nitrogen (N) status already during the first rotation period; (2) input of symbiotic fixation covers an essential part of the plant annual N demand of the stand; (3) despite a considerable N input into the ecosystem of a young alder stand, there will occur no significant environmental hazards (N leaching or N₂O emissions). The first two hypotheses can be accepted: there was a significant increase in N and C content in the topsoil (from 0.11 to 0.14%, and from 1.4 to 1.7%, respectively), and N fixation (151.5 kg N ha⁻¹ yr⁻¹) covered about 74% of the annual N demand of the stand. The third hypothesis met support as well: N₂O emissions (0.5 kg N ha⁻¹ yr⁻¹) were low, while most of the annual gaseous N losses were in the form of N₂ (73.8 kg N ha⁻¹ yr⁻¹). Annual average NO₃-N leaching was 15 kg N ha⁻¹ yr⁻¹ but the N that leached from topsoil accumulated in deeper soil layers. The soil acidifying effect of alders was clearly evident; during the 14-year period soil acidity increased 1.3 units in the upper 0-10 cm topsoil layer.     

Fine root classification matters: nutrient levels in different functional categories,orders and diameters of roots in boreal Pinus sylvestris across a latitudinal gradient

Plant and Soil - Any grouping of tree species concerned with SOC sequestration should include trees that are as homogeneous as possible in their carbon sequestration. We propose a classification of...     

Does the age of fine root carbon indicate the age of fine roots in boreal forests?

To test the reliability of the radiocarbon method for determining root age, we analyzed fine roots (originating from the years 1985?C1993) from ingrowth cores with known maximum root age (1?C6?years old). For this purpose, three Scots pine (Pinus sylvestris L.) stands were selected from boreal forests in Finland. We analyzed root ¹⁴C age by the radiocarbon method and compared it with the above-mentioned known maximum fine root age. In general, ages determined by the two methods (root ¹⁴C age and ingrowth core root maximum age) were in agreement with each other for roots of small diameter (<0.5?mm). By contrast, in most of the samples of fine roots of larger diameter (1.5?C2?mm), the ¹⁴C age of root samples of 1987?C1989 exceeded the ingrowth core root maximum age by 1?C10?years. This shows that these roots had received a large amount of older stored carbon from unknown sources in addition to atmospheric CO₂ directly from photosynthesis. We conclude that the ¹⁴C signature of fine roots, especially those of larger diameter, may not always be indicative of root age, and that further studies are needed concerning the extent of possible root uptake of older carbon and its residence time in roots.     

Variation of carbon age of fine roots in boreal forests determined from 14C measurements

Background and aims

The main objectives of this study were to determine how the carbon age of fine root cellulose varies between stands, tree species, root diameter and soil depth. In addition, we also compared the carbon age of fine roots from soil cores of this study with reported values from the roots of the same diameter classes of ingrowth cores on the same sites.

Methods

We used natural abundance of ¹⁴C to estimate root carbon age in four boreal Norway spruce and Scots pine stands in Finland and Estonia.

Results

Age of fine root carbon was older in 1.5–2 mm diameter fine roots than in fine roots with <0.5 mm diameter, and tended to be older in mineral soil than in organic soil. Fine root carbon was older in the less fertile Finnish spruce stands (11–12 years) than in the more fertile Estonian stand (3 and 8 years), implying that roots may live longer in less fertile soil. We further observed that on one of our sites carbon in live fine roots with the 1.5–2 mm diameter was of similar C age (7–12 years) than in the ingrowth core roots despite the reported root age in the ingrowth cores – being not older than 2 years.

Conclusions

From this result, we conclude that new live roots may in some cases use old carbon reserves for their cellulose formation. Future research should be oriented towards improving our understanding of possible internal redistribution and uptake of C in trees.     

Fine-root growth,mortality and heavy metal concentrations in limed and fertilized Pinus silvestris (L.) stands in the vicinity of a Cu-Ni smelter in SW Finland

This study was conducted to assess 1) the growth of fine roots into ingrowth cores and fine root mortality, 2) the effects of liming and correction fertilization on fine-root growth and mortality, and 3) the concentrations of heavy metals in fine roots in control, limed or fertilized Scots pine stands at different distances from a copper-nickel smelter. Fine-root biomass in the ingrowth cores in the control plots varied between 1 (at 0.5 km from the smelter) and 252 and 271 g/m² (at 4 and 8 km, respectively). In the most polluted stand at 0.5 km, 98% of the fine roots that had grown into the ingrowth cores had died before sampling. Corresponding values for the other stands (4 and 8 km) were only 13-18%. At 0.5 km, liming increased the growth and survival of fine roots. The concentrations of Cu and Ni were also smaller in fine roots from the limed plot than those from the control plot. In the correction fertilization treatment at 0.5 km the total ingrowth of fine roots was at the same level as in the control, but less fine roots had died. Thus, the correction fertilizer treatment increased the survival but not the growth of fine roots. At 4 or 8 km, there were no significant differences in the fine-root biomass or necromass or element concentrations between the treatments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

Fine root turnover and litter production of Norway spruce in a long-term temperature and nutrient manipulation experiment

Background and aims

Increased soil temperature and nutrient availability enhance soil biological activity. We studied how these affect fine root growth and survival, i.e. below-ground litter production, in relation to above-ground foliage litter production of Norway spruce (Picea abies (L.) Karst.).

Methods

The treatments, irrigation (I), soil warming + irrigation (WI), fertilization + irrigation (FI) and soil warming + fertilization + irrigation (WFI) were started in 1987 (F, I) and in 1995 (W). The annual production of fine root litter was estimated from minirhizotrons (survival) and soil-cores (biomass) and the annual above-ground litter production from litter traps.

Results and conclusions

The number and elongation of fine roots tended to be higher in WI and I compared to the other treatments, which may indicate nutrient shortage. Fine roots in the WFI treatment had the lowest median longevity and from three to fourfold higher below-ground litter production compared to WI, FI or I - higher soil temperature increased the litter input particularly into the mineral soil. Only fertilization increased the above-ground litter production. As warmer and more nutrient-rich soil significantly shortened the fine root lifespan and increased the litter input, the storage of carbon in boreal forest soil may increase in the future.     

Effects of long-term temperature and nutrient manipulation on Norway spruce fine roots and mycelia production

Aims and methods

The effects of changing climate on ectomycorrhizal (EcM) fine roots were studied in northern Sweden by manipulating soil temperature for 14 years and/or by fertilizing for 22 years. Fine root biomass, necromass, EcM root tip biomass, morphology and number as well as mycelia production were determined from soil cores and mesh bags.

Results and conclusions

The fine root biomass and necromass were highest in the fertilized plots, following similar trends in the above-ground biomass, whereas the EcM root tip biomass per basal area decreased by 22 % in the fertilized plots compared to the control. Warming increased the fine root biomass, live/dead-ratio and the number of EcM root tips in the mineral soil and tended to increase the production of EcM mycelia. Greater fine root biomass meant more EcM root tips, although the tip frequency was not affected by fertilization or warming. Significantly higher specific root length of EcM root tips indicated an increased need for nutrients in warmed and in unfertilized plots. Better nutrient supply and warmer soil temperature provide a potential to increase the flow of carbon into the soil via increased fine root biomass, but the carbon balance also depends on root turnover.