The age of fine-root carbon in three forests of the eastern United States measured by radiocarbon

Using a new approach involving one-time measurements of radiocarbon (¹⁴C) in fine (<2 mm diameter) root tissues we have directly measured the mean age of fine-root carbon. We find that the carbon making up the standing stock of fine roots in deciduous and coniferous forests of the eastern United States has a mean age of 3-18 years for live fine roots, 10-18 years for dead fine roots, and 3-18 years for mixed live+dead fine roots. These ¹⁴C-derived mean ages represent the time C was stored in the plant before being allocated for root growth, plus the average lifespan (for live roots), plus the average time for the root to decompose (for dead roots and mixtures). Comparison of the ¹⁴C content of roots known to have grown within 1 year with the ¹⁴C of atmospheric CO₂ for the same period shows that root tissues are derived from recently fixed carbon, and the storage time prior to allocation is <2 years and likely <1 year. Fine-root mean ages tend to increase with depth in the soil. Live roots in the organic horizons are made of C fixed 3-8 years ago compared with 11-18 years in the mineral B horizons. The mean age of C in roots increases with root diameter and also is related to branching order. Our results differ dramatically from previous estimates of fine-root mean ages made using mass balance approaches and root-viewing cameras, which generally report life spans (mean ages for live roots) of a few months to 1-2 years. Each method for estimating fine-root dynamics, including this new radiocarbon method, has biases. Root-viewing approaches tend to emphasize more rapidly cycling roots, while radiocarbon ages tend to reflect those components that persist longest in the soil. Our ¹⁴C-derived estimates of long mean ages can be reconciled with faster estimates only if fine-root populations have varying rates of root mortality and decomposition. Our results indicate that a standard definition of fine roots, as those with diameters of <2 mm, is inadequate to determine the most dynamic portion of the root population. Recognition of the variability in fine-root dynamics is necessary to obtain better estimates of belowground C inputs.     

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.     

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.     

Partitioning sources of soil-respired CO2 and their seasonal variation using a unique radiocarbon tracer

Soil respiration is derived from heterotrophic (decomposition of soil organic matter) and autotrophic (root/rhizosphere respiration) sources, but there is considerable uncertainty about what factors control variations in their relative contributions in space and time. We took advantage of a unique whole‐ecosystem radiocarbon label in a temperate forest to partition soil respiration into three sources: (1) recently photosynthesized carbon (C), which dominates root and rhizosphere respiration; (2) leaf litter decomposition and (3) decomposition of root litter and soil organic matter >1–2 years old. Heterotrophic sources and specifically leaf litter decomposition were large contributors to total soil respiration during the growing season. Relative contributions from leaf litter decomposition ranged from a low of ～1±3% of total soil respiration (6± 3 mg C m^?2 h^?1) when leaf litter was extremely dry, to a high of 42±16% (96± 38 mg C m^?2 h^?1). Total soil respiration fluxes varied with the strength of the leaf litter decomposition source, indicating that moisture‐dependent changes in litter decomposition drive variability in total soil respiration fluxes. In the surface mineral soil layer, decomposition of C fixed in the original labeling event (3–5 years earlier) dominated the isotopic signature of heterotrophic respiration. Root/rhizosphere respiration accounted for 16±10% to 64±22% of total soil respiration, with highest relative contributions coinciding with low overall soil respiration fluxes. In contrast to leaf litter decomposition, root respiration fluxes did not exhibit marked temporal variation ranging from 34±14 to 40±16 mg C m^?2 h^?1 at different times in the growing season with a single exception (88±35 mg C m^?2 h^?1). Radiocarbon signatures of root respired CO₂ changed markedly between early and late spring (March vs. May), suggesting a switch from stored nonstructural carbohydrate sources to more recent photosynthetic products.     

Partitioning sources of soil respiration in boreal black spruce forest using radiocarbon

Separating ecosystem and soil respiration into autotrophic and heterotrophic component sources is necessary for understanding how the net ecosystem exchange of carbon (C) will respond to current and future changes in climate and vegetation. Here, we use an isotope mass balance method based on radiocarbon to partition respiration sources in three mature black spruce forest stands in Alaska. Radiocarbon (Δ¹⁴C) signatures of respired C reflect the age of substrate C and can be used to differentiate source pools within ecosystems. Recently‐fixed C that fuels plant or microbial metabolism has Δ¹⁴C values close to that of current atmospheric CO₂, while C respired from litter and soil organic matter decomposition will reflect the longer residence time of C in plant and soil C pools. Contrary to our expectations, the Δ¹⁴C of C respired by recently excised black spruce roots averaged 14‰ greater than expected for recently fixed photosynthetic products, indicating that some portion of the C fueling root metabolism was derived from C storage pools with turnover times of at least several years. The Δ¹⁴C values of C respired by heterotrophs in laboratory incubations of soil organic matter averaged 60‰ higher than the contemporary atmosphere Δ¹⁴CO₂, indicating that the major contributors to decomposition are derived from a combination of sources consistent with a mean residence time of up to a decade. Comparing autotrophic and heterotrophic Δ¹⁴C end members with measurements of the Δ¹⁴C of total soil respiration, we calculated that 47–63% of soil CO₂ emissions were derived from heterotrophic respiration across all three sites. Our limited temporal sampling also observed no significant differences in the partitioning of soil respiration in the early season compared with the late season. Future work is needed to address the reasons for high Δ¹⁴C values in root respiration and issues of whether this method fully captures the contribution of rhizosphere respiration.     

模拟氮沉降对杉木幼苗细根的生理生态影响

细根对氮沉降的生理生态响应将显著影响森林生态系统的生产力和碳吸存。为了揭示氮沉降对杉木细根的生理生态影响,对一年生杉木(Cunninghamia lanceolata)幼苗进行了模拟氮沉降试验,并测定施氮1年后杉木幼苗细根生物量、细根形态学特征(比根长、比表面积)、元素化学计量学指标(C、N、P、C/N、C/P、N/P)、细根代谢特征(细根比呼吸速率、非结构性碳水化合物)。结果表明:(1)杉木细根生物量随氮添加水平的升高而显著降低,尤其是0—1 mm细根生物量;细根比根长和比表面积随氮添加水平升高而显著增大。(2)氮添加后杉木细根C含量、C/N、C/P显著降低,高氮添加导致1—2 mm细根N含量和N/P显著升高,而低氮添加导致1—2 mm细根P含量显著升高、N/P显著降低,而0—1 mm细根的N、P含量则保持相对稳定。(3)氮添加后杉木细根比呼吸速率无显著变化,细根可溶性糖含量随氮添加增加而显著增加,而淀粉含量和NSC显著降低。综合以上结果表明:氮添加后用于细根形态构建的碳分配减少,这可能会减少土壤中有机碳的保留,0—1 mm细根的形态更易发生变化,但是其内部N、P养分含量相对更稳定以维持生理活动,细根NSC对氮添加的响应表明施氮可能导致细根受光合产物的限制。     

Fine-root turnover patterns and their relationship to root diameter and soil depth in a 14C-labeled hardwood forest

Characterization of turnover times of fine roots is essential to understanding patterns of carbon allocation in plants and describing forest C cycling. We used the rate of decline in the ratio of 14C to 12C in a mature hardwood forest, enriched by an inadvertent 14C pulse, to investigate fine-root turnover and its relationship with fine-root diameter and soil depth. Biomass and Delta14C values were determined for fine roots collected during three consecutive winters from four sites, by depth, diameter size classes (< 0.5 or 0.5-2 mm), and live-or-dead status. Live-root pools retained significant 14C enrichment over 3 yr, demonstrating a mean turnover time on the order of years. However, elevated Delta14C values in dead-root pools within 18 months of the pulse indicated an additional component of live roots with short turnover times (months). Our results challenge assumptions of a single live fine-root pool with a unimodal and normal age distribution. Live fine roots < 0.5 mm and those near the surface, especially those in the O horizon, had more rapid turnover than 0.5-2 mm roots and deeper roots, respectively.     

Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings

In this study, we investigated the impact of elevated atmospheric CO₂ (ambient + 350 μmol mol^–1) on fine root production and respiration in Scots pine (Pinus sylvestris L.) seedlings. After six months exposure to elevated CO₂, root production measured by root in-growth bags, showed significant increases in mean total root length and biomass, which were more than 100% greater compared to the ambient treatment. This increased root length may have lead to a more intensive soil exploration. Chemical analysis of the roots showed that the roots in the elevated treatment accumulated more starch and had a lower C/N-ratio. Specific root respiration rates were significantly higher in the elevated treatment and this was probably attributed to increased nitrogen concentrations in the roots. Rhizospheric respiration and soil CO₂ efflux were also enhanced in the elevated treatment. These results clearly indicate that under elevated atmospheric CO₂ root production and development in Scots pine seedlings is altered and respiratory carbon losses through the root system are increased.     

Changing sources of soil respiration with time since fire in a boreal forest

Radiocarbon signatures (Δ¹⁴C) of carbon dioxide (CO₂) provide a measure of the age of C being decomposed by microbes or respired by living plants. Over a 2‐year period, we measured Δ¹⁴C of soil respiration and soil CO₂ in boreal forest sites in Canada, which varied primarily in the amount of time since the last stand‐replacing fire. Comparing bulk respiration Δ¹⁴C with Δ¹⁴C of CO₂ evolved in incubations of heterotrophic (decomposing organic horizons) and autotrophic (root and moss) components allowed us to estimate the relative contributions of O horizon decomposition vs. plant sources. Although soil respiration fluxes did not vary greatly, differences in Δ¹⁴C of respired CO₂ indicated marked variation in respiration sources in space and time. The ¹⁴C signature of respired CO₂ respired from O horizon decomposition depended on the age of C substrates. These varied with time since fire, but consistently had Δ¹⁴C greater (averaging ～120‰) than autotrophic respiration. The Δ¹⁴C of autotrophically respired CO₂ in young stands equaled those expected for recent photosynthetic products (70‰ in 2003, 64‰ in 2004). CO₂ respired by black spruce roots in stands >40 years old had Δ¹⁴C up to 30‰ higher than recent photosynthates, indicating a significant contribution of C stored at least several years in plants. Decomposition of O horizon organic matter made up 20% or less of soil respiration in the younger (<40 years since fire) stands, increasing to ～50% in mature stands. This is a minimum for total heterotrophic contribution, since mineral soil CO₂ had Δ¹⁴C close to or less than those we have assigned to autotrophic respiration. Decomposition of old organic matter in mineral soils clearly contributed to soil respiration in younger stands in 2003, a very dry year, when Δ¹⁴C of soil respiration in younger successional stands dropped below those of the atmospheric CO₂.     

Size and Structure of Fine Root Systems in Old-growth and Secondary Tropical Montane Forests (Costa Rica)

The fine root systems of three tropical montane forests differing in age and history were investigated in the Cordillera Talamanca, Costa Rica. We analyzed abundance, vertical distribution, and morphology of fine roots in an early successional forest (10–15 years old, ESF), a mid‐successional forest (40 years old, MSP), and a nearby undisturbed old‐growth forest (OGF), and related the root data to soil morphological and chemical parameters. The OGF stand contained a 19 cm deep organic layer on the forest floor (i.e., 530 mol C/m²), which was two and five times thicker than that of the MSF (10 cm) and ESF stands (4 cm), respectively. There was a corresponding decrease in fine root biomass in this horizon from 1128 g dry matter/m² in the old‐growth forest to 337 (MSF) and 31 g/m² (ESF) in the secondary forests, although the stands had similar leaf areas. The organic layer was a preferred substrate for fine root growth in the old‐growth forest as indicated by more than four times higher fine root densities (root mass per soil volume) than in the mineral topsoil (0–10 cm); in the two secondary forests, root densities in the organic layer were equal to or lower than in the mineral soil. Specific fine root surface areas and specific root tip abundance (tips per unit root dry mass) were significantly greater in the roots of the ESF than the MSF and OGF stands. Most roots of the ESF trees (8 abundant species) were infected by VA mycorrhizal fungi; ectomycorrhizal species (Quercus copeyemis and Q. costaricensis) were dominant in the MSF and OGF stands. Replacement of tropical montane oak forest by secondary forest in Costa Rica has resulted in (1) a large reduction of tree fine root biomass; (2) a substantial decrease in depth of the organic layer (and thus in preferred rooting space); and (3) a great loss of soil carbon and nutrients. Whether old–growth Quercus forests maintain a very high fine root biomass because their ectomycorrhizal rootlets are less effective in nutrient absorption than those of VA mycorrhizal secondary forests, or if their nutrient demand is much higher than that of secondary forests (despite a similar leaf area and leaf mass production), remains unclear.     

Fine root dynamics and trace gas fluxes in two lowland tropical forest soils

Fine root dynamics have the potential to contribute significantly to ecosystem‐scale biogeochemical cycling, including the production and emission of greenhouse gases. This is particularly true in tropical forests which are often characterized as having large fine root biomass and rapid rates of root production and decomposition. We examined patterns in fine root dynamics on two soil types in a lowland moist Amazonian forest, and determined the effect of root decay on rates of C and N trace gas fluxes. Root production averaged 229 (±35) and 153 (±27) g m^?2 yr^?1 for years 1 and 2 of the study, respectively, and did not vary significantly with soil texture. Root decay was sensitive to soil texture with faster rates in the clay soil (k=?0.96 year^?1) than in the sandy loam soil (k=?0.61 year^?1), leading to greater standing stocks of dead roots in the sandy loam. Rates of nitrous oxide (N₂O) emissions were significantly greater in the clay soil (13±1 ng N cm^?2 h^?1) than in the sandy loam (1.4±0.2 ng N cm^?2 h^?1). Root mortality and decay following trenching doubled rates of N₂O emissions in the clay and tripled them in sandy loam over a 1‐year period. Trenching also increased nitric oxide fluxes, which were greater in the sandy loam than in the clay. We used trenching (clay only) and a mass balance approach to estimate the root contribution to soil respiration. In clay soil root respiration was 264–380 g C m^?2 yr^?1, accounting for 24% to 35% of the total soil CO₂ efflux. Estimates were similar using both approaches. In sandy loam, root respiration rates were slightly higher and more variable (521±206 g C m² yr^?1) and contributed 35% of the total soil respiration. Our results show that soil heterotrophs strongly dominate soil respiration in this forest, regardless of soil texture. Our results also suggest that fine root mortality and decomposition associated with disturbance and land‐use change can contribute significantly to increased rates of nitrogen trace gas emissions.     

Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2-enrichment: a six-year-minirhizotron study

Efforts to characterize carbon (C) cycling among atmosphere, forest canopy, and soil C pools are hindered by poorly quantified fine root dynamics. We characterized the influence of free‐air‐CO₂‐enrichment (ambient +200 ppm) on fine roots for a period of 6 years (Autumn 1998 through Autumn 2004) in an 18‐year‐old loblolly pine (Pinus taeda) plantation near Durham, NC, USA using minirhizotrons. Root production and mortality were synchronous processes that peaked most years during spring and early summer. Seasonality of fine root production and mortality was not influenced by atmospheric CO₂ availability. Averaged over all 6 years of the study, CO₂ enrichment increased average fine root standing crop (+23%), annual root length production (+25%), and annual root length mortality (+36%). Larger increase in mortality compared with production with CO₂ enrichment is explained by shorter average fine root lifespans in elevated plots (500 days) compared with controls (574 days). The effects of CO₂‐enrichment on fine root proliferation tended to shift from shallow (0–15 cm) to deeper soil depths (15–30) with increasing duration of the study. Diameters of fine roots were initially increased by CO₂‐enrichment but this effect diminished over time. Averaged over 6 years, annual fine root NPP was estimated to be 163 g dw m^?2 yr^?1 in CO₂‐enriched plots and 130 g dw m^?2 yr^?1 in control plots (P= 0.13) corresponding to an average annual additional input of fine root biomass to soil of 33 g m^?2 yr^?1 in CO₂‐enriched plots. A lack of consistent CO₂× year effects suggest that the positive effects of CO₂ enrichment on fine root growth persisted 6 years following minirhizotron tube installation (8 years following initiation of the CO₂ fumigation). Although CO₂‐enrichment contributed to extra flow of C into soil in this experiment, the magnitude of the effect was small suggesting only modest potential for fine root processes to directly contribute to soil C storage in south‐eastern pine forests.     

Contribution of Lolium perenne rhizodeposition to carbon turnover of pasture soil

Carbon rhizodeposition and root respiration during eight development stages of Lolium perenne were studied on a loamy Gleyic Cambisol by ¹⁴CO₂ pulse labelling of shoots in a two compartment chamber under controlled laboratory conditions. Total ¹⁴CO₂ efflux from the soil (root respiration, microbial respiration of exudates and dead roots) in the first 8 days after ¹⁴C pulse labelling decreased during plant development from 14 to 6.5% of the total ¹⁴C input. Root respiration accounted for was between 1.5 and 6.5% while microbial respiration of easily available rhizodeposits and dead root remains were between 2 and 8% of the ¹⁴C input. Both respiration processes were found to decline during plant development, but only the decrease in root respiration was significant. The average contribution of root respiration to total ¹⁴CO₂ efflux from the soil was approximately 41%. Close correlation was found between cumulative ¹⁴CO₂ efflux from the soil and the time when maximum ¹⁴CO₂ efflux occurred (r=0.97). The average total of CO₂ Defflux from the soil with Lolium perenne was approximately 21 μg C-CO₂ d⁻¹ g⁻¹. It increased slightly during plant development. The contribution of plant roots to total CO₂ efflux from the soil, calculated as the remainder from respiration of bare soil, was about 51%. The total ¹⁴C content after 8 days in the soil with roots ranged from 8.2 to 27.7% of assimilated carbon. This corresponds to an underground carbon transfer by Lolium perenne of 6–10 g C m⁻² at the beginning of the growth period and 50–65 g C m⁻² towards the end of the growth period. The conventional root washing procedure was found to be inadequate for the determination of total carbon input in the soil because 90% of the young fine roots can be lost. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date. This revised version was published online in June 2006 with corrections to the Cover Date.     

An in situ approach for measuring root-associated respiration and nitrate uptake of forest trees

The ecophysiological characteristics of fine roots of mature forest plants are poorly understood because of difficulties of measurement. We explored a root in-growth approach to measure respiration and nitrate uptake of woody plant roots in situ. Roots of seven species were grown into sand-filled chambers. Root-associated respiration was measured as CO ₂ emission on four dates and nitrate uptake was quantified using ¹⁵N. All the roots were younger than 3 months at the time of measurement. Fine root respiration measured over the temperature range of 14.5–15.5 °C averaged 18.9–36.5 nmol gDM ^–1 s ^–1 across species. Nitrate uptake rates by these fine roots (1.3–6.8 nmol gDM ^–1 s ^–1) were comparable to other studies of forest trees. The root respiration rates were several times higher than measurements on detached roots of mature trees, concurring with literature observations that young roots respire much more rapidly than older roots. The root in-growth approach appears promising for providing information on the metabolic activity of fine roots of mature forest trees growing in soil.     

Clean rain promotes fine root growth and soil respiration in a Norway spruce forest

Soil acidification and N saturation are considered to affect the decomposition of soil organic matter as well as growth and mortality of fine roots in many forest soils. Here we report from a field experiment where ‘clean rain’ has been applied to the soil for about 10 years under a roofed plot of a 71‐year‐old Norway spruce plantation at Solling, Central Germany. Reduced amounts of protons (?78%), sulphate (?53%), ammonium (?86%), and nitrate (?49%) were sprayed on the soil surface of the clean rain plot between 1992 and 2001. In an adjacent roofed control plot, throughfall was collected and immediately re‐sprinkled below the roof construction without any chemical manipulation. One year before the clean rain treatment started, live and dead fine root masses (≤2 mm) were determined from undisturbed soil cores down to 40 cm mineral soil depth. Total live fine root mass was significantly lower in the clean rain plot than in the control plot. After the first sampling, the soil holes were refilled with quartz sand and repeatedly sampled in June 1992, June 1996, and October 2001. There were no differences in live and dead fine root masses between the plots in 1992 and 1996. In 2001, both live and dead fine root masses of the clean rain plot were about twice as high as in the control plot, indicating that fine root growth recovered in the mineral soil following 10 years of clean rain treatment. Moreover, the clean rain treatment significantly reduced the total N concentrations of live fine roots and 1‐year‐old needles. Our results suggest that the reduced N input promoted fine root growth to compensate N deficiency. Reduced Al concentration in soil solution may have contributed to the recovery of fine root growth, however, the toxicity of Al species is largely unknown. Mean annual soil respiration rate was 24% higher in the period from 2000 to 2001, indicating that the clean rain treatment increased respiration of roots and heterotrophic microorganisms within the rhizosphere. Laboratory incubation of samples from the organic horizon and the top mineral soil revealed no differences between the plots in the decay rate of soil organic matter. Our results suggest that strong reductions in atmospheric N deposition from about 30 to 10 kg N ha^?1 yr^?1 and decreasing acid stress can have beneficial effects on growth of fine roots in the mineral soil within a decade. We conclude that biological recovery under reduced atmospheric loads can affect the nutrient and carbon budget of spruce soils in the long run.     

Effects of elevated atmospheric CO2 on fine root production and activity in an intact temperate forest ecosystem

We investigated the effects of elevated atmospheric CO₂ concentrations (ambient + 200 ppm) on fine root production and soil carbon dynamics in a loblolly pine (Pinus taeda) forest subject to free‐air CO₂ enrichment (FACE) near Durham, NC (USA). Live fine root mass (LFR) showed less seasonal variation than dead fine root mass (DFR), which was correlated with seasonal changes in soil moisture and soil temperature. LFR mass increased significantly (by 86%) in the elevated CO₂ treatment, with an increment of 37 g(dry weight) m^?2 above the control plots after two years of CO₂ fumigation. There was no long‐term increment in DFR associated with elevated CO₂, but significant seasonal accumulations of DFR mass occurred during the summer of the second year of fumigation. Overall, root net primary production (RNPP) was not significantly different, but annual carbon inputs were 21.7 gC m^?2 y^?1 (68%) higher in the elevated CO₂ treatment compared to controls. Specific root respiration was not altered by the CO₂ treatment during most of the year; however, it was significantly higher by 21% and 13% in September 1997 and May 1998, respectively, in elevated CO₂. We did not find statistically significant differences in the C/N ratio of the root tissue, root decomposition or phosphatase activity in soil and roots associated with the treatment. Our data show that the early response of a loblolly pine forest ecosystem subject to CO₂ enrichment is an increase in its fine root population and a trend towards higher total RNPP after two years of CO₂ fumigation.     

Does low soil base saturation affect fine root properties of European beech (Fagus sylvatica L.)?

It is generally believed that high soil solution Al³⁺ in acidic soils with low base saturation (BS), negatively influences the properties of fine roots. Fine roots from European beech (Fagus sylvatica L.) trees growing in highly acidic soils with very low BS and potentially high Al³⁺ concentration in the soil solution were analysed and the dependency of fine root properties on soil BS was measured. The fine roots were sampled down to 1 m depth at seven forest sites located on the Swiss Plateau. These sites varied in their BS from 1.4 to 11.4% in the mineral layers. We evaluated relationships between the BS of these mineral layers and fine root properties, such as ratio between bio- and necromass (live/dead ratio), specific root length (SRL), root tip abundance (RTA), root branching abundance (RBA), O₂-consumption, and the Ca/Al molar ratio in the fine root tissue. The fine root properties were compared not only with the BS of the soil, but also with the Ca/Al molar ratio in the fine root tissues. Significant relations of fine root properties occurred when the soils of the seven sites were grouped into two BS groups (<5 and 5–10%). The live/dead ratio, the RTA, the RBA, the O₂-consumption, and Ca/Al molar ratio were lower in the group of BS <5% than in the group 5–10%. Decreases in the morphological properties and in the O₂-consumption were related to decrease in the Ca/Al molar ratio of the fine root tissues. There is evidence that the fine root properties are negatively influenced, nevertheless, fine root systems of mature European beech in their natural ecological environment seem to be able to compensate adverse effects of low BS. Responsible Editor: Philippe Hinsinger.     

Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.     

Belowground drought response of European beech: fine root biomass and carbon partitioning in 14 mature stands across a precipitation gradient

How tree root systems will respond to increased drought stress, as predicted for parts of Central Europe, is not well understood. According to the optimal partitioning theory, plants should enhance root growth relative to aboveground growth in order to reduce water limitations. We tested this prediction in a transect study with 14 mature forest stands of European beech (Fagus sylvatica L.) by analysing the response of the fine root system to a large decrease in annual precipitation (970–520 mm yr⁻¹). In 3 years with contrasting precipitation regimes, we investigated leaf area and leaf biomass, fine root biomass and necromass (organic layer and mineral soil to 40 cm) and fine root productivity (ingrowth core approach), and analysed the dependence on precipitation, temperature, soil nutrient availability and stand structure. In contrast to the optimal partitioning theory, fine root biomass decreased by about a third from stands with >950 mm yr⁻¹ to those with <550 mm yr⁻¹, while leaf biomass remained constant, resulting in a significant decrease, and not an increase, in the fine root/leaf biomass ratio towards drier sites. Average fine root diameter decreased towards the drier stands, thereby partly compensating for the loss in root biomass and surface area. Both δ¹³C‐signature of fine root mass and the ingrowth core data indicated a higher fine root turnover in the drier stands. Principal components analyses (PCA) and regression analyses revealed a positive influence of precipitation on the profile total of fine root biomass in the 14 stands and a negative one of temperature and plant‐available soil phosphorus. We hypothesize that summer droughts lead to increased fine root mortality, thereby reducing root biomass, but they also stimulate compensatory fine root production in the drier stands. We conclude that the optimal partitioning theory fails to explain the observed decrease in the fine root/leaf biomass ratio, but is supported by the data if carbon allocation to roots is considered, which would account for enhanced root turnover in drier environments.