Fine-root dynamics in sugi (Cryptomeria japonica) under manipulated soil nitrogen conditions

Aims

Nitrogen deposition affect fine-root dynamics, a key factor in forest carbon and nutrient dynamics. This study aimed to elucidate the effects of increased soil inorganic nitrogen (N) levels on the fine-root dynamics of Cryptomeria japonica, which is tolerant to excess N load.

Methods

An ammonium nitrate solution (28 kg ha^?1 month^?1) was applied for 3 years to plots (1 m?×?2 m) in a C. japonica plantation. The elongation and disappearance of the fine roots were examined using the minirhizotron technique.

Results

The N fertilization increased soil inorganic N content and lowered the soil pH. Fine-root elongation rates increased with fertilization, whereas patterns of their seasonal changes were not affected. The ratio of cumulative disappearance to cumulative elongation of fine roots was lower in the N-fertilized plots than in the control plots. The mean diameter of the fine roots was not affected by N fertilization.

Conclusions

Our results suggest that C. japonica can respond to increased levels of soil inorganic N by increasing both the production and residence time of the fine roots. However, the effects of the changing soil N content are less evident for the phenology and morphology of the fine roots in C. japonica.     

Effects of litter types, microsite and root diameters on litter decomposition in Pinus sylvestris plantations of northern China

Background and aims

Litter decomposition is a major process in the carbon (C) flow and nutrient cycling of terrestrial ecosystems, but the effects of litter type, microsite, and root diameter on decomposition are poorly understood.

Methods

Litterbags were used to examine the decomposition rate of leaf litter and roots at three soil depths (5, 10 and 20 cm) over a 470-day period in Pinus sylvestris plantations in northern China.

Results

Leaves and the finest roots decomposed more quickly at 5 cm depth and coarser roots (>1-mm) decomposed more quickly at 10 and 20 cm depth. Roots generally decomposed more quickly than leaf litter, except at 5 cm deep; leaves decomposed more quickly than the coarsest roots (>5-mm). Root decomposition was strongly influenced by root diameter. Leaves experienced net nitrogen (N) immobilization and coarse roots (>2-mm) experienced more N release than fine roots. Significant heterogeneity was seen in N release for fine-roots (<2-mm) with N immobilization occurring in smaller (0.5–2-mm) roots and N release in the finest roots (<0.5-mm).

Conclusions

Soil depth of litter placement significantly influenced the relative contribution of the decomposition of leaves and roots of different diameters to carbon and nutrient cycling.     

Fine-root trait plasticity of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>) and spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) forests on two contrasting soils

Aim

The fine roots of trees may show plastic responses to their resource environment. Several, contrasting hypotheses exist on this plasticity, but empirical evidence for these hypotheses is scattered. This study aims to enhance our understanding of tree root plasticity by examining intra-specific variation in fine-root mass and morphology, fine-root growth and decomposition, and associated mycorrhizal interactions in beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) forests on soils that differ in resource availability.

Methods

We measured the mass and morphological traits of fine roots (i.e. ≤ 2 mm diameter) sampled to 50 cm depth. Fine-root growth was measured with ingrowth cores, and fine-root decomposition with litter bags. Mycorrhizal fungal biomass was determined using ingrowth mesh bags.

Results

Both tree species showed more than three times higher fine-root mass, and a ten-fold higher fine-root growth rate on sand than on clay, but no or marginal differences in overall fine-root morphology. Within the fine-root category however, beech stands had relatively more root length of their finest roots on clay than on sand. In the spruce stands, ectomycorrhizal mycelium biomass was larger on sand than on clay.

Conclusions

In temperate beech and spruce forests, fine-root mass and mycorrhizal fungal biomass, rather than fine-root morphology, are changed to ensure uptake under different soil resource conditions. Yet enhancing our mechanistic understanding of fine-root trait plasticity and how it affects tree growth requires more attention to fine-root dynamics, the functional diversity within the fine-roots, and mycorrhizal symbiosis as an important belowground uptake strategy.

     

Slow decomposition and limited nitrogen release by lower order roots in eight Chinese temperate and subtropical trees

Background and aims

Roots of the lowest branch orders have the highest mortality rate, and may contribute predominately to plant carbon (C) and nutrient transfer into the soil. Yet patterns and controlling factors of the decomposition of these roots are poorly understood.

Methods

We conducted a two-year field litterbag study on different root orders and leaf litter in four temperate and four subtropical tree species.

Results

Five species showed slower decay rates in lower- (order 1–2) than higher-order (order 3–5) roots, and all species showed slower decay rates in lower-order roots than leaf litter. These patterns were strongly related to higher acid-insoluble fraction in lower- than higher-order roots, and in roots than in leaf litter, but were unrelated to initial N concentration. Litter N was predominantly in recalcitrant forms and limited amount of N was released during the study period;only 12 % of root N and 26 % of leaf litter N was released in 2 years.

Conclusions

We conclude that the slow decomposition of lower-order roots may be a common phenomenon and is mainly driven by their high acid-insoluble fraction. Moreover, litter N, especially root N, is retained during decomposition and may not be available for immediate plant uptake.     

Variation in the aboveground stand structure and fine-root biomass of Bornean heath (kerangas) forests in relation to altitude and soil nitrogen availability

Key message

Borneo’s tropical heath (kerangas) forest has limited soil nutrient availability, and high variation in aboveground structure and fine-root biomass. This variation depends on altitude and soil nitrogen availability.

Abstract

To elucidate the biotic and abiotic factors affecting the variation in fine-root biomass (FRB, <2 mm diameter) of trees growing under nutrient-poor environments in Sabah, North Borneo, we investigated FRB in different forests with varying soil nitrogen (N) availability. We selected two study sites at different altitudes: the Maliau Basin (ca. 1000 m asl) and Nabawan (ca. 500 m asl). Both sites included tropical heath (kerangas) forest, on infertile soils (podzols) with a surface organic horizon overlying a bleached (eluviated) mineral horizon, and taller forests on more fertile non-podzolic soils. FRB was obtained from each plot by soil coring (to a depth of 15 cm). FRB increased with decreasing soil inorganic N content (NH₄–N and NO₃–N), tree height, and aboveground biomass. Thus, higher proportions of carbon resources were allocated to fine-roots in stands with lower N availability. FRB was significantly greater at the Maliau Basin than at Nabawan, reflecting lower soil N availability at higher altitude. Our results demonstrate high variation in FRB among the heath forests, and suggest that fine-root development is more prominent under a cooler climate where N availability limits tree growth owing to slower decomposition. The variation in N availability under the same climate (i.e., at the same altitude) appears to be related to the extent of soil podzolization.

     

Relation of fine root distribution to soil C in a Cunninghamia lanceolata plantation in subtropical China

Background and aims

Growth and distribution of fine roots closely depend on soil resource availability and affect soil C distribution in return. Understanding of relationships between fine root distribution and soil C can help to predict the contribution of fine root turnover to soil C accumulation.

Methods

A study was conducted in a subtropical Cunninghamia lanceolata plantation to assess the fine root mass density (FRMD), fine root C density (FRCD) of different fine root groups as well as their relations with soil C.

Results

The FRMD and FRCD of short-lived roots, dead roots and herb roots peaked in the 0–10 cm soil layer and decreased with soil depth, while FRMD, FRCD of long-lived roots peaked in the 10–20 cm soil layer. Soil C was positively related to FRMD and FRCD of total fine roots (across all three soil layers), dead roots (0–10 cm) and herb roots (10–20 cm) as well as FRCD of short-lived roots (20–40 cm) (P <0.05).

Conclusions

Soil C was mainly affected by herb roots in upper soil layers and by woody plant roots in deeper soil layers.     

Root dynamics of Carex stricta-dominated tussock meadows

Background and aims

The roots of tussock-forming plants contribute to the formation of microtopographic features in many ecosystems, but the dynamics of such roots are poorly understood. We examined the spatial heterogeneity of tussock fine root dynamics to investigate allocation patterns and the role of root productivity in the persistence of tussock structures.

Methods

We compared the spatial variability of fine root (<1 mm, 1–2 mm) density, biomass, % live, allocation, turnover rate (using bomb 14C), and productivity of four Carex stricta Lam.-dominated tussock meadows in the upper Midwest, USA (3 reference, 1 restored site).

Results

Relative to underlying microsites, tussocks were warm, dry, and high in root density, productivity, % live biomass, and turnover. Root productivity averaged 649 g?m^?2 yr^?1 (±208) in reference sites, comprised 57 % (±10) of total net production, and was concentrated in tussocks (70 %?±?4). Root turnover rate averaged 0.63 yr^?1 (±0.08), but tussocks had ~50 % faster root turnover than the underlying soil, and <1 mm roots turned over ~40 % faster than 1–2 mm roots.

Conclusions

Our detailed analysis of the spatial heterogeneity of tussock root dynamics suggests that high allocation and elevated turnover of tussock roots facilitates organic matter accumulation and tussock persistence over time.     

Root distribution of different mature tree species growing on contrasting textured soils in temperate windbreaks

Aim

We studied the vertical and lateral root distribution of tree species from three genera (Populus spp. - poplar, Picea spp. - spruce, Salix spp. - willow) that were planted in temperate windbreaks and assessed the effects of soil texture on root density.

Methods

Root distribution to depths of up to 1 m was assessed using the trench-profile method at different distances from the tree rows (2, 6 and 9 m) in 18 mature (average age, 25 years-old) windbreak-sites that were located on light- or heavy-textured agricultural soils in southeastern Québec, Canada. Roots were classified into three diameter classes: fine (<1 mm), medium-size (1–5 mm), and coarse (>5 mm).

Results

Tree fine-root density in poplar and willow windbreaks was higher than in spruce windbreaks at 2 m from the tree row. Root densities were higher in light compared to heavy soils, but these differences were specific to poplar and spruce. Across species groups and soil types, 67 % of the roots occurred in the uppermost 30 cm. In this soil zone, different soil fertility variables (pH, clay content, CEC) were negatively correlated with root density. Densities of spruce and willow roots at 6 m from the tree row were much lower (and often unobserved) than that of poplar. At 9 m, low root densities were observed at only two sites.

Conclusions

We conclude that tree identity and soil type are important drivers of root distribution in temperate agroforestry systems. These results may have important implications for the management of tree competition in agroforestry systems and several ecosystem services that are provided by roots, including C-sequestration, erosion control and water infiltration.     

Tree species mixture inhibits soil organic carbon mineralization accompanied by decreased r-selected bacteria

Background and aims

Fine-root functioning is a major driver of plant growth and strongly influences the global carbon cycle. While fine-root over-yielding has been shown in the upper soil layers of mixed-species forests relative to monospecific stands, the consequences of tree diversity on fine-root growth in very deep soil layers is still unknown. Our study aimed to assess the consequences of mixing Acacia mangium and Eucalyptus grandis trees on soil exploration by roots down to the water table at 17 m depth in a tropical planted forest.

Method

Fine roots (diameter < 2 mm) were sampled in a randomized block design with three treatments: monospecific stands of Acacia mangium (100A), Eucalyptus grandis (100E), and mixed stands with 50% of each species (50A50E). Root ingrowth bags were installed at 4 depths (from 0.1 m to 6 m) in the three treatments within three different blocks, to study the fine-root production over 2 periods of 3 months.

Results

Down to 17 m depth, total fine-root biomass was 1127 g m^?2 in 50A50E, 780 g m^?2 in 100A and 714 g m^?2 in 100E. Specific root length and specific root area were 110–150% higher in 50A50E than in 100A for Acacia mangium trees and 34% higher in 50A50E than in 100E for Eucalyptus grandis trees. Ingrowth bags showed that the capacity of fine roots to explore soil patches did not decrease down to a depth of 6 m for the two species.

Conclusions

Belowground interactions between Acacia mangium and Eucalyptus grandis trees greatly increased the exploration of very deep soil layers by fine roots, which is likely to enhance the uptake of soil resources. Mixing tree species might therefore increase the resilience of tropical planted forests through a better exploration of deep soils.

     

The measurement of woody root decomposition using two methodologies in a Sitka spruce forest ecosystem

Background and aims

The decomposition of roots is an important process in the loss of carbon (C) and the mineralization of nitrogen (N) in forest ecosystems. The early stage decomposition rate of Sitka spruce (Picea sitchensis (Bong.) Carr.) roots was determined using trenched plots and decomposition bags.

Methods

Stumps of known age were trenched and quadrants (50?cm by 50?cm) excavated from randomly selected stumps every 6?months over 4?years, while the mass loss from buried roots in decomposition bags, divided among four diameter categories (ranging from fine roots <2?mm to large roots >50?mm), was monitored for 27?months. The C and N concentrations of excavated samples at different time points were analysed.

Results

The change in total root necromass per quadrant showed a higher decomposition rate-constant (k) of 0.24?±?0.068?year^?1 than the k-value of roots in decomposition bags (0.07?±?0.005?year^?1). The C concentration (47.24?±?0.609?%) did not significantly change with decomposition. There was a significant increase in the C:N ratio of roots in all diameter categories (fine: 48.92?%, small: 38.53?%, medium 11.71?%, large: 76.25?%) after 4?years of decomposition, driven by N loss. Root diameter accounted for 78?% of the variation in the N concentration of roots as decomposition progressed.

Conclusion

Though the trenched plot approach offered an alternative to the more common decomposition bag method for estimating root decomposition, high spatial variation and sampling difficulties may lead to an overestimation of the mass loss from trenched roots, thus, the decomposition bag method gives a more reliable decomposition rate-constant.     

Factors controlling decomposition rates of fine root litter in temperate forests and grasslands

Background and aims

Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 μm mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors.

Results

Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24?±?6 % mass loss after 12 months, mean ± SD) than beech roots in forest soils (12?±?4 %; p?Conclusions Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality.     

Fine root biomass in two black spruce stands in interior Alaska: effects of different permafrost conditions

Key message

In black spruce stands on permafrost, trees and understory plants showed higher biomass allocation especially to ‘thin’ fine roots (diam. < 0.5 mm) when growing on shallower permafrost table.

Abstract

Black spruce (Picea mariana) forests in interior Alaska are located on permafrost and show greater below-ground biomass allocation than non-permafrost forests. However, information on fine roots (roots <2 mm in diameter), which have a key role in nutrient uptake and below-ground carbon flux, is still limited especially for effects of different permafrost conditions. In this study, we examined fine root biomass in two black spruce stands with different depths to the permafrost table. In the shallow permafrost (SP) plot, fine root biomass of black spruce trees was 70 % of that in the deep permafrost (DP) plot. In contrast, ratio of the fine root biomass to above-ground biomass was greater in the SP plot than in the DP plot. Understory plants, on the other hand, showed larger fine root biomass in the SP plot than in the DP plot, whereas their above-ground biomass was similar between the two plots. In addition, biomass proportion of ‘thin’ fine roots (diam. <0.5 mm) in total fine roots was greater in the SP plot than in the DP plot. These results suggest that black spruce trees and understory plants could increase biomass allocation to fine roots for efficient below-ground resource acquisition from colder environments with shallower permafrost table. In the SP plot, fine roots of understory plants accounted for 30 % of the stand fine root biomass, suggesting that understory plants such as Ledum and Vaccinium spp. would have significant contribution to below-ground carbon dynamics in permafrost forests.

     

Root decomposition of <Emphasis Type="Italic">Artemisia halodendron</Emphasis> and its effect on soil nitrogen and soil organic carbon in the Horqin Sandy Land,northeastern China

Root decomposition is a critical feedback from the plant to the soil, especially in sandy land where strong winds remove aboveground litter. As a pioneer shrub in semi-mobile dunes of the Horqin sandy land, Artemisia halodendron has multiple effects on nutrient capture and the microenvironment. However, its root decomposition has not been studied in terms of its influence on soil organic carbon (SOC) and nitrogen (N). In this study, we buried fine (≤2 mm) and coarse roots in litterbags at a depth of 15 cm below semi-mobile dunes. We measured the masses remaining and the C and N contents at intervals during 434 days of decomposition. The soils below the litterbags were then divided into layers and sampled to measure the SOC and N contents. After rapid initial decomposition, both coarse and fine roots decomposed slowly. After 53 days, 36.2 % of coarse roots and 39.8 % of fine roots had decomposed. In contrast, only 18.4 % of coarse roots and 30.5 % of fine roots decomposed in the following 381 days. Fine roots decomposed significantly faster, and their decomposition rate after the initial rapid decay was strongly related to climate (R ² = 0.716, P < 0.05). Root decomposition increased SOC and N contents below the litterbags, with larger effects for fine roots. The SOC content was more variable between soil layers than the N content. Thus, decomposition of A. halodendron roots cannot be ignored when studying SOC and N feedbacks from plants to the soil, particularly for fine roots.     

Variety specific relationships between effects of rhizobacteria on root exudation,growth and nutrient uptake of soybean

Aims

It has been increasingly recognized that only distal lower order roots turn over actively within the <2 mm fine root system of trees. This study aimed to estimate fine root production and turnover rate based on lower order fine roots and their relations to soil variables in mangroves.

Methods

We conducted sequential coring in five natural mangrove forests at Dongzhai Bay, China. Annual fine root production and turnover rate were calculated based on the seasonal variations of the biomass and necromass of lower order roots or the whole fine root system.

Results

Annual fine root production and turnover rate ranged between 571 and 2838 g m^?2 and 1.46–5.96 yr^?1, respectively, estimated with lower order roots, and they were increased by 0–30 % and reduced by 13–48 %, respectively, estimated with the whole fine root system. Annual fine root production was 1–3.5 times higher than aboveground litter production and was positively related to soil carbon, nitrogen and phosphorus concentrations. Fine root turnover rate was negatively related to soil salinity.

Conclusions

Mangrove fine root turnover plays a more important role than aboveground litter production in soil C accumulation. Sites with higher soil nutrients and lower salinity favor fine root production and turnover, and thus favor soil C accumulation.

     

Improved scaling of minirhizotron data using an empirically-derived depth of field and correcting for the underestimation of root diameters

Background and aims

Accurate data on the standing crop, production, and turnover of fine roots is essential to our understanding of major terrestrial ecological processes. Minirhizotrons offer a unique opportunity to study the dynamic processes of root systems, but are susceptible to several measurement biases.

Methods

We use roots extracted from minirhizotron tube surfaces to calculate the depth of field of a minirhizotron image and present a model to correct for the underestimation of root diameters obscured by soil in minirhizotron images.

Results

Non-linear regression analysis resulted in an estimated depth of field of 0.78 mm for minirhizotron images. Unadjusted minirhizotron data underestimated root net primary production and fine root standing crop by 61 % when compared to adjusted data using our depth of field and root diameter corrections. Changes in depth of field accounted for >99 % of standing crop adjustments with root diameter corrections accounting for <1 %.

Conclusions

Our results represent the first effort to empirically derive depth of field for minirhizotron images. This work may explain the commonly reported underestimation of fine roots using minirhizotrons, and stands to improve the ability of researchers to accurately scale minirhizotron data to large soil volumes.     

An optimized fine root sampling methodology balancing accuracy and time investment

Aims

Tree roots are spatially highly heterogeneous and it thus requires large numbers of samples to detect statistically significant changes in root biomass. The objectives of this study were to understand and quantify the sources of error in the assessment of fine root biomass (<2 mm) during the second year of a high-density Populus plantation.

Methods

Soil cores were collected in winter (n?=?35) and in summer (n?=?20), and fine roots were picked by hand for varying lengths of time: 1, 2, 5, 20, 40, and 60 min. The root biomass data were used to identify the best combination of the time spent for root picking and the number of samples collected, that minimizes the overall uncertainty (i.e. the combination of the spatial error due to the incomplete sampling and the temporal error due to the incomplete core processing).

Results

On average, 25 min was enough time to pick 90 % of the fine root biomass in winter, while in summer only 10 min were needed. In winter fewer samples were needed, but more time for picking was necessary as compared to summer when root biomass was higher.

Conclusions

Fine root sampling can be optimized by minimizing the uncertainty of the biomass estimates and simultaneously decreasing root sampling time investment.     

Fine root dynamics and soil carbon accretion under thinned and un-thinned Cupressus lusitanica stands in, Southern Ethiopia

Background and aims

Forest management activities influences stand nutrient budgets, belowground carbon allocation and storage in the soil. A field experiment was carried out in Southern Ethiopia to investigate the effect of thinning on fine root dynamics and associated soil carbon accretion of 6-year old C. lusitanica stands.

Methods

Fine roots (≤2 mm in diameter) were sampled seasonally to a depth of 40 cm using sequential root coring method. Fine root biomass and necromass, vertical distribution, seasonal dynamics, annual turnover and soil carbon accretion were quantified.

Results

Fine root biomass and necromass showed vertical and temporal variations. More than 70 % of the fine root mass was concentrated in the top 20 cm soil depth. Fine root biomass showed significant seasonal variation with peaks at the end of the major rainy season and short rainy season. Thinning significantly increased fine root necromass, annual fine root production and turnover. Mean annual soil carbon accretion, through fine root necromass, in the thinned stand was 63 % higher than that in the un-thinned stand.

Conclusions

The temporal dynamics in fine roots is driven by the seasonality in precipitation. Thinning of C. lusitanica plantation would increase soil C accretion considerably through increased fine root necromass and turnover.     

Fine root dry matter relative to mango (<Emphasis Type="Italic">Mangifera indica</Emphasis>) tree scion size grafted on size-controlling rootstocks,is negatively related to scion growth rate

Key message

The effects of mango rootstock cultivars on scion vigour may be predicted by scion growth rate being negatively related to fine root dry matter/scion trunk cross sectional area.

Abstract

Knowledge of root dry matter (DM) allocation, in relation to differing vigour conferred by rootstock cultivars, is required to understand the structural relationships between rootstock and scion. We investigated the mass of roots (four size classes up to 23 mm diameter) by coring proximal to five polyembryonic mango rootstock cultivars known to differ in their effects on the vigour and productivity of scion cultivar ‘Kensington Pride’, in a field trial of 13-year-old trees. Significant differences in fine (<0.64 and 0.64–1.88 mm diameter) and small (1.88–7.50 mm) root DM contents were observed between rootstock cultivars. There was a complex relationship between the amount of feeder (fine and small size classes) roots and scion size (trunk cross sectional area, TCSA), with intermediate size trees on rootstock MYP having the most feeder roots, while the smallest trees, on the rootstock Vellaikulamban had the least of these roots. Across rootstock cultivars, tree vigour (TCSA growth rate) was negatively and significantly related to the ratio of fine root DM/scion TCSA, suggesting this may be a useful indicator of the vigour that different rootstocks confer on the scion. In contrast non-ratio root DM and scion TCSA results had no significant relationships. The significant rootstock effects on orchard root growth and tree size could not be predicted from earlier differences in nursery seedling vigour, nor did seedling vigour predict root DM allocation.

     

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.