The role of soil conditions in fine root ecomorphology in Norway spruce (Picea abies (L.) Karst.)

The present study is an attempt to investigate the pattern of morphological variability of the short roots of Norway spruce (Picea abies (L.) Karst.) growing in different soils. Five root parameters – diameter, length and dry weight of the root tip, root density (dry weight per water-saturated volume) and specific root area (absorbing area of dry weight unit) were studied with respect to 11 soil characteristics using CANOCO RDA analysis. The investigation was conducted in seven study areas in Estonia differing in site quality class and soil type. Ten root samples per study area were collected randomly from the forest floor and from the 20 cm soil surface layer. Eleven soil parameters were included in the study: humus content, specific soil surface area, field capacity, soil bulk density, pH (KCl and H₂O dilution's), N and Ca concentrations, Ca/Al and C/N ratios, and the decomposition rate of fine roots (<2 mm dia.). Root morphological characteristics most strongly related to the measured soil characteristics in the different sites were specific root area, root density and diameter of the short roots, the means varying from 29 to 42 m² kg⁻¹, from 310 to 540 kg m⁻³ and from 0.26 to 0.32 mm, respectively; root density being most sensitive. The most favourable site and soil types resulting in fine roots with morphological characteristics for optimizing nutrient uptake (e.g. low short root density and high specific root area) were Umbric Luvisol (Oxalis), Dystric Gleysol (Oxalis) and Gleyic Luvisol (Hepatica). These soil types correspond to highly productive natural forest stands of Norway spruce in Estonia. All measured soil variables explained 28% of total variance of the root characteristics. The most important variables related to root morphology were the humus content, field capacity and specific soil surface area. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Root turnover in a beech and a spruce stand of the Belgian Ardennes

The theoretical basis of fine root turnover estimation in forest soils is discussed, in relation to appropriate experimental techniques of measurement. After sequential coring, the correct expression is the sum of significant positive increments of live and dead roots of the various diameter categories, to which the transfer of dead roots to organic matter derived from roots, OMDR, has to be added. This should not be confounded with dead root mineralization. The transfer rates should first be estimated in root dimensions and not in weight of dry matter. The measurements were carried out in a 120 year old beech (Fagus sylvatica L.) stand and a 35 year old Norway spruce (Picea abies Karst) stand, in the Eastern Ardennes, Belgium. The turnover rate of fine roots (diam. <5 mm) was 4393 kg ha⁻¹ year⁻¹ (root dry weight), including 711.2 kg ha⁻¹ year⁻¹ for dead root transfer to OMDR, for beech. For spruce, turnover rate was 7011 kg ha⁻¹ year⁻¹ (root dry weight), including 1498 kg ha⁻¹ year⁻¹ for dead root transfer to OMDR. Under beech, there was a slight root density increase in spring. No seasonal fluctuations were observed under spruce, but a strong irreversible drop in live root growth was found in the later season 1980–1981, corresponding to a decrease of tree height growth and trunk radius increment. Turnover rates were further expressed in dry weight and in amounts of elements (kg ha⁻¹ year⁻¹) (Ca, Mg, K, Na, Al, N, P, S). Correlative relations between root dimensions and dry weight and element concentrations show that the derived values, and in particular root specific density (dry weight volume⁻¹) vary according to species, root category, and seasonal sampling. Various schemes of seasonal variations of root growth, described in Europe, show that the major dependance on general climate is obscured by environmental factors (soil, exposure, species). It is suggested that root density fluctuation approach the steady state on an annual basis under mild Atlantic conditions.     

Specific root area: A soil characteristic

The root systems of all plants adapt to soil conditions. One of the ways Norway spruce adapts is a regulation of absorbing root surface area, which can be viewed as an ecomorphological index of soil conditions. Minimal specific root surface area of absorbing roots of Norway spruce (28–29 m²kg⁻¹) responds to optimal soil conditions. This was achieved at a soil bulk density of about 1.2 gcm⁻³.     

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.     

Fine root vertical distribution and temporal dynamics in mature stands of two enset (Enset ventricosum Welw Cheesman) clones

Quantification of the role of fine roots in the biological cycle of nutrients necessitates understanding root distribution, estimating root biomass, turnover rate and nutrient concentrations, and the dynamics of these parameters in perennial systems. Temporal dynamics, vertical distribution, annual production and turnover, and nitrogen use of fine roots (≤2 mm in diameter) were studied in mature (5-year-old) stands of two enset (Ensete ventricosum) clones using the in-growth bag technique. Live fine root mass generally decreased with increasing depth across all seasons except the dry period. Except for the dry period, more than 70% of the fine root mass was in the above 0-20 cm depth, and the fine root mass in the upper 0–10 cm depth was significantly higher than in the lowest depth (20–30 cm). Live fine root mass showed a seasonal peak at the end of the major rainy season but fell to its lowest value during the dry or short rainy season. The difference between the peak and low periods were significant (p ≤ 0.05). Fine root nitrogen (N) use showed significant seasonal variation where the mean monthly fine root N use was highest during the major rainy season. There were significant effects on N use due to depths and in-growth periods, but not due to clones. Enset fine root production and turnover ranged from 2,339 to 2,451 kg ha⁻¹ year⁻¹ and from 1.55 to 1.80 year⁻¹, respectively. Root N return, calculated from fine root turnover, was estimated at 64–65 kg ha⁻¹ year⁻¹. Fine root production, vertical distribution and temporal dynamics may be related to moisture variations and nutrient (N) fluxes among seasons and along the soil depth. The study showed that fine root production and turnover can contribute considerably to the carbon and nitrogen economy of mature enset plots.     

Effects of elevated soil solution Al concentrations on fine roots in a middle-aged Norway spruce (Picea abies (L.) Karst.) stand

Aluminium (Al), mobilized by acidic deposition, has been claimed to be a major threat to forest vitality. Fine root mortality, decreased root growth and reduced nutrient uptake have been observed in controlled laboratory experiments where roots of tree seedlings were exposed to elevated concentrations of Al. Yet, evidence for Al-induced root damage from forest stands is scarcely reported. Nevertheless, Al dissolved in soil water has received a key role in the critical load concept for forests. Here, we present effects of artificially elevated concentrations of Al in the soil solution on fine roots in a middle-aged stand of Norway spruce (Picea abies (L.) Karst.). Although the inorganic Al concentrations about 200 µM and Ca:Al ratio about 0.7 that were established in the soil solution within this experiment have been associated with reduction of root growth and root mortality for spruce seedlings in hydroponic studies, no acute damage on fine roots was observed. Three years of treatment did not cause visual root damage, nor were effects on fine root necromass observed. Fine root necromass made up about 10% of fine root biomass for all treatments. However, significantly lower molar Ca:Al and Mg:Al ratios in living and dead fine roots were found in the plots where Al concentrations were highest and ratios of Ca to Al in the soil solution were lowest. The lack of response on fine root biomass suggests that forest stands tolerate higher Al levels than results from laboratory experiments indicate. We conclude that effect studies in the laboratory have limited value for field conditions. The key role of Al toxicity, expressed as the Ca/Al ratio, in critical load calculations for forests may have to be reconsidered.     

Effect of artificial soil acidification and liming on growth and nutrient status of mycorrhizal roots of Norway spruce (Picea abies [L.] Karst.)

Effects of soil acidification and liming on biomass responses and free Al, Ca, K, Mg, Mn and P contents of mycorrhizal roots of mature Norway spruce (Picea abies [L.] Karst.) were studied at Höglwald Forest in Southern Germany.At the untreated site, mycorrhizal root biomass was lower in the acid humus (pH = 3.3) than in the less acid upper (0–5 cm) mineral soil (pH 4.1). Mycorrhizal roots from the humus contained 10% of the level free Al in mycorrhizal roots from the upper mineral soil. During seven years of soil acidification the quantity of mycorrhizal roots remained unaffected in the humus and the upper mineral soil, perhaps due to the high buffering capacity of the humus which prevented a significant alteration of the nutrient status of the roots. However, two years after soil acidification had been terminated, the percentage of mycorrhizal roots in the humus decreased, possibly because the free root concentrations of K had decreased.On the other hand, six years after liming, there was a two-fold increase of the annual mean quantity of mycorrhizal roots in the humus. Compensatory liming (acid irrigation after liming) had a similar effect on mycorrhizal root production in the humus. However, two years after acid irrigation had been terminated a decrease of mycorrhizal roots in the upper mineral soil (0–5 cm) was observed. Since the total amount of mycorrhizal roots in the humus and upper mineral soil remained constant, compensatory liming produced a shift in fine roots to the humus layer.The higher mass of living mycorrhizal roots in the upper mineral soil (0–5 cm) as compared to the humus of the untreated plot as well as the increased mass of mycorrhizal roots in the humus after liming or compensatory liming are both attributed to an increase in pH to 4.5 rather than alleviation of Al toxicity.     

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.     

Influence of cultivation method on root grafting in Norway spruce (Picea abies (L.) Karst.)

Root grafting is the process by which a functional union of two or more roots subsequent to their formation is formed. The above- and below-ground parts of three Norway spruce stands (natural stand, Umbric Luvisol; row-culture and group-culture, Planosol; stand ages 40, 43 and 43, respectively) of high site quality (I) were investigated. Stand densities were 1550, 1783 and 1722 stems ha^-1, respectively. In all investigated stands, root grafting was most sensitive to tree spacing. Grafts were observed in case the distance between the trees was 0.7–1.2 m. Grafts occurred always in areas of higher rooting density, in a row of the row-culture and within a tree group in the group-culture. Root grafting was enhanced in case of a narrower humus horizon in the group culture compared with the row-culture, 16.5 and 30 cm, respectively; the humus horizon contained 99% and 95% of conducting roots with d ≥ 5 mm, respectively. Root graftings occurred in 75% of excavated trees in the group-culture, in 37.5% of excavated trees in the row-culture and in 33.3% of excavated trees in the natural stand. Stand age was 24 years in the row-culture and 22 years in the group-culture at the beginning of root grafting. No grafts occurred between two suppressed trees, whereas in 86–100% of all cases, at least one tree was dominant or codominant. In row- and group-cultivated Norway spruce stands, the initial minimum diameter of the grafted root without bark was from 1 to 3 cm in 63% of cases. Grafting of roots with d < 1 cm or d > 10 cm was rare or absent. Root grafting had usually begun at the root age of 10–20 years (46% of cases). This revised version was published online in June 2006 with corrections to the Cover Date.     

Wood-ash recycling affects forest soil and tree fine-root chemistry and reverses soil acidification

Wood ash was applied to a forest ecosystem with the aim to recycle nutrients taken from the forest and to mitigate the negative effects of intensive harvesting. After two years, the application of 8,000 kg ha⁻¹ of wood ash increased soil exchangeable Ca and Mg. Similarly, an increase in Ca and Mg in the Norway spruce fine roots was recorded, leading to significant linear correlations between soil and root Ca and soil and root Mg. In contrast to these macronutrients, the micronutrients Fe and Zn and the toxic element Al decreased in the soil exchangeable fraction with the addition of wood ash, but not in the fine roots. Only Mn decreased in soil and in fine roots leading to a significant linear correlation between soil and root Mn. In soil, as well as in fine roots, strong positive correlations were found between the elements Ca and Mg and between Fe and Al. This indicates that the uptake of Mg resembles that of Ca and that of Al that of Fe. With the wood ash application, the pH increased from 3.2 to 4.8, the base saturation from 30% to 86%, the molar basic cations/Al ratio (BC/Al) of the soil solution from 1.5 to 5.5, and the molar Ca/Al ratio of the fine roots from 1.3 to 3.7. Overall, all below-ground indicators of soil acidification responded positively to the wood ash application within two years. Nitrate concentrations increased only slightly in the soil solution at a soil depth of 75–80 cm, and no signs of increased heavy metal concentrations in the soils or in the fine roots were apparent. This suggests that the recycling of wood ash could be an integral part of sustainable forest management because it closes the nutrient cycle and reverses soil acidification.     

Distribution of roots and root nodules and biomass allocation in young intensively managed grey alder stands on a peat bog

Development of below-ground biomass and biomass allocation were studied in two different stands of young grey alder stands growing on a peat bog. Both stands were given the same fertilization and irrigation treatment. The roots were investigated from 1) open plastic tubes enclosing the complete root systems in 1982, and 2) root cores 1984–86. Coarse roots (diameter>1 mm) were mainly found close to the trunk of the trees while fine roots (≤1 mm) were more evenly distributed in the stands. Root nodules were intermediate in distribution. The root systems were shallow, with more than 90% of the biomass in the uppermost 9–10 cm of the soil, probably because of low oxygen availability in the peat soil. The biomass allocation to the above-ground parts increased during the study period.     

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root and needle litter decomposition responses to enhanced supplies of N and S in a Norway spruce forest in southwest Sweden

The effects of additions of ammonium sulfate (NS) on the decomposition of litter derived from Norway spruce roots (< 2 and 2 - 5 mm in diameter) in the humus and mineral soil layers (0 - 15 cm) of a Norway spruce stand in southern Sweden were investigated over a 6-year period. To this purpose, litterbags were incubated in the humus layer and in the mineral soil in June 1996, with roots collected from NS and control (C) plots incubated in the NS and C plots, respectively. The N concentrations in fine roots (< 2 mm) in the NS- plots were higher than those in 2 - 5 mm roots in both humus and mineral soil layers. In the humus layer, N concentrations in the fine roots in the C- and NS- plots were 12.8 and 15.7 mg g ^{? 1}, respectively. By the end of the fifth year the < 2 mm roots in humus layer had lost 48.5 and 50% of their mass in the C and NS plots, respectively, while the corresponding values for the 2 - 5 mm diameter class were 44 and 54%. The fresh root litter may be a sensitive indicator to responses to enhanced N and S deposition, although decomposition rates of both litter types are affected.     

落叶松人工林细根动态与土壤资源有效性关系研究

树木细根在森林生态系统C和养分循环中具有重要的作用。由于温带土壤资源有效性具有明显的季节变化, 导致细根生物量、根长密度 (Rootlengthdensity, RLD) 和比根长 (Specificrootlength, SRL) 的季节性变化。以 17年生落叶松 (Larixgmelini) 人工林为研究对象, 采用根钻法从 5月到 10月连续取样, 研究了不同土层细根 (直径≤ 2mm) 生物量、RLD和SRL的季节动态, 以及这些根系指标动态与土壤水分、温度和N有效性的关系。结果表明 :1) 落叶松细根年平均生物量 (活根 +死根 ) 为 189.1g·m-2 ·a-1, 其中 5 0 %分布在表层 (0~ 10cm), 33%分布在亚表层 (11~ 2 0cm), 17%分布在底层 (2 1~ 30cm) 。活根和死根生物量在 5~ 7月以及 9月较高, 8月和 10月较低。从春季 (5月 ) 到秋季 (10月 ), 随着活细根生物量的减少, 死细根生物量增加 ;2 ) 土壤表层 (0~ 10cm) 具有较高的RLD和SRL, 而底层 (2 1~ 30cm) 最低。春季 (5月 ) 总RLD和SRL最高, 分别为 10 6 2 1.4 5m·m-3 和 14.83m·g-1, 到秋季 (9月 ) 树木生长结束后达到最低值, 分别为 2 198.2 0m·m-3 和 3.77m·g-1;3) 细根生物量、RLD和SRL与土壤水分、温度和有效N存在不同程度的相关性。从单因子分析来看, 土壤水分和有效N对细根的影响明显大于温度, 对活根的影响大于死根。由于土壤资源有效性的季节变化, 使得C的地下分配格局发生改变。各土层细根与有效性资源之间的相关性反映了细根功能季节性差异。细根 (生物量、RLD和SRL) 的季节动态 (5 8%~ 73%的变异 ) 主要由土壤资源有效性的季节变化引起。     

Nitrogen dynamics of a boreal black spruce wildfire chronosequence

This study examined the nitrogen (N) dynamics of a black spruce (Picea mariana (Mill.) BSP)-dominated chronosequence in Manitoba, Canada. The seven sites studied each contained separate well- and poorly drained stands, originated from stand-killing wildfires, and were between 3 and 151 years old. Our goals were to (i) measure total N concentration ([N]) of all biomass components and major soil horizons; (ii) compare N content and select vegetation N cycle processes among the stands; and (iii) examine relationships between ecosystem C and N cycling for these stands. Vegetation [N] varied significantly by tissue type, species, soil drainage, and stand age; woody debris [N] increased with decay state and decreased with debris size. Soil [N] declined with horizon depth but did not vary with stand age. Total (live + dead) biomass N content ranged from 18.4 to 99.7 g N m⁻² in the well-drained stands and 37.8–154.6 g N m⁻² in the poorly drained stands. Mean soil N content (380.6 g N m⁻²) was unaffected by stand age. Annual vegetation N requirement (5.9 and 8.4 g N m⁻² yr⁻¹ in the middle-aged well- and poorly drained stands, respectively) was dominated by trees and fine roots in the well-drained stands, and bryophytes in the poorly drained stands. Fraction N retranslocated was significantly higher in deciduous than evergreen tree species, and in older than younger stands. Nitrogen use efficiency (NUE) was significantly lower in bryophytes than in trees, and in deciduous than in evergreen trees. Tree NUE increased with stand age, but overall stand NUE was roughly constant (∼ ∼150 g g⁻¹ N) across the entire chronosequence.     

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.     

Measuring Fine Root Turnover in Forest Ecosystems

Development of direct and indirect methods for measuring root turnover and the status of knowledge on fine root turnover in forest ecosystems are discussed. While soil and ingrowth cores give estimates of standing root biomass and relative growth, respectively, minirhizotrons provide estimates of median root longevity (turnover time) i.e., the time by which 50% of the roots are dead. Advanced minirhizotron and carbon tracer studies combined with demographic statistical methods and new models hold the promise of improving our fundamental understanding of the factors controlling root turnover. Using minirhizotron data, fine root turnover (y⁻¹) can be estimated in two ways: as the ratio of annual root length production to average live root length observed and as the inverse of median root longevity. Fine root production and mortality can be estimated by combining data from minirhizotrons and soil cores, provided that these data are based on roots of the same diameter class (e.g., < 1 mm in diameter) and changes in the same time steps. Fluxes of carbon and nutrients via fine root mortality can then be estimated by multiplying the amount of carbon and nutrients in fine root biomass by fine root turnover. It is suggested that the minirhizotron method is suitable for estimating median fine root longevity. In comparison to the minirhizotron method, the radio carbon technique favor larger fine roots that are less dynamics. We need to reconcile and improve both methods to develop a more complete understanding of root turnover.