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.     

Soil fertility and fine root dynamics in response to 4 years of nutrient (N,P, K) fertilization in a lowland tropical moist forest,Panama

The question of how tropical trees cope with infertile soils has been challenging to address, in part, because fine root dynamics must be studied in situ. We used annual fertilization with nitrogen (N as urea, 12.5 g N m^?2 year^?1), phosphorus (P as superphosphate, 5 g P m^?2 year^?1) and potassium (K as KCl, 5 g K m^?2 year^?1) within 38 ha of old‐growth lowland tropical moist forest in Panama and examined fine root dynamics with minirhizotron images. We expected that added P, above all, would (i) decrease fine root biomass but, (ii) have no impact on fine root turnover. Soil in the study area was moderately acidic (pH = 5.28), had moderate concentrations of exchangeable base cations (13.4 cmol kg^?1), low concentrations of Bray‐extractable phosphate (PO₄ = 2.2 mg kg^?1), and modest concentrations of KCl‐extractable nitrate (NO₃ = 5.0 mg kg^?1) and KCl‐extractable ammonium (NH₄ = 15.5 mg kg^?1). Added N increased concentrations of KCl‐extractable NO₃ and acidified the soil by one pH unit. Added P increased concentrations of Bray‐extractable PO₄ and P in the labile fraction. Concentrations of exchangeable K were elevated in K addition plots but reduced by N additions. Fine root dynamics responded to added K rather than added P. After 2 years, added K decreased fine root biomass from 330 to 275 g m^?2. The turnover coefficient of fine roots <1 mm diameter ranged from 2.6 to 4.4 per year, and the largest values occurred in plots with added K. This study supported the view that biomass and dynamics of fine roots respond to soil nutrient availability in species‐rich, lowland tropical moist forest. However, K rather than P elicited root responses. Fine roots smaller than 1 mm have a short lifetime (<140 days), and control of fine root production by nutrient availability in tropical forests deserves more study.     

Nutrient resorption responses to water and nitrogen amendment in semi-arid grassland of Inner Mongolia, China

Litterfall and fine root production is a major pathway for carbon and nutrient cycling in forest ecosystems. We investigated leaf litterfall, fine-root mass, production and turnover rate in the upper soil (0–30 cm) under four major tree species (Leucaena leucocephala, Acacia nilotica, Azadirachta indica, Prosopis juliflora) of the semi-arid region of India. All the four tree species showed an unimodal peak of leaf litterfall with distinct seasonality. Leucaena leucocephala and Acacia nilotica had maximum leaf litterfall between September and December while Azadirachta indica and Prosopis juliflora shed most of their leaves between February and May. Annual leaf litterfall of the four species ranged from 3.3 Mg ha^?1 (Leucaena leucocephala) to 8.1 Mg ha^?1 (Prosopis juliflora). Marked seasonal variations in amount of fine root biomass were observed in all the four tree species. Fine root production was maximum in Prosopis juliflora (171 g m^?2 y^?1) followed by Azadirachta indica (169 g m^?2 y^?1), Acacia nilotica (106 g m^?2 y^?1) and Leucaena leucocephala (79 g m^?2 y^?1). Fine root biomass showed a seasonal peak after the rainy season but fell to its lowest value during the winter and dry summer season. Fine root turnover rate ranged from 0.56 to 0.97 y^?1 and followed the order Azadirachta indica > Leucaena leucocephala > Prosopis juliflora > Acacia nilotica. The results of this study demonstrated that Prosopis juliflora and Azadirachta indica had greater capability for maintaining site productivity as evidenced from greater leaf litterfall and fine root production.     

Fine root growth phenology,production, and turnover in a northern hardwood forest ecosystem

A large part of the nutrient flux in deciduous forests is through fine root turnover, yet this process is seldom measured. As part of a nutrient cycling study, fine root dynamics were studied for two years at Huntington Forest in the Adirondack Mountain region of New York, USA. Root growth phenology was characterized using field rhizotrons, three methods were used to estimate fine root production, two methods were used to estimate fine root mortality, and decomposition was estimated using the buried bag technique. During both 1986 and 1987, fine root elongation began in early April, peaked during July and August, and nearly ceased by mid-October. Mean fine root ( 3 mm diameter) biomass in the surface 28-cm was 2.5 t ha^–1 and necromass was 2.9 t ha^–1. Annual decomposition rates ranged from 17 to 30% beneath the litter and 27 to 52% at a depth of 10 cm. Depending on the method used for estimation, fine root production ranged from 2.0 to 2.9 t ha^–1, mortality ranged from 1.8 to 3.7 t ha^–1 yr^–1, and decomposition was 0.9 t ha^–1 yr^–1. Thus, turnover ranged from 0.8 to 1.2 yr^–1. The nutrients that cycled through fine roots annually were 4.5–6.1 kg Ca, 1.1–1.4 kg Mg, 0.3–0.4 kg K, 1.2–1.7 kg P, 20.3–27.3 kg N, and 1.8–2.4 kg S ha^–1. Fine root turnover was less important than leaf litterfall in the cycling of Ca and Mg and was similar to leaf litterfall in the amount of N, P, K and S cycled.     

Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above‐ and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha^?1 at 194 m to 4.95 ± 0.62 Mg C ha^?1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha^?1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha^?1 at 194 m to 47.03 Mg C ha^?1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha^?1 yr^?1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha^?1 yr^?1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha^?1 yr^?1 (210 m) and 2.59 ± 0.40 Mg C ha^?1 yr^?1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above‐ and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Fine root turnover of irrigated hedgerow intercropping in Northern Kenya

Fine root turnover (<2 mm) was determined from repeated measurements of root distribution up to 120 cm soil depth by core sampling in four month intervals. Sole cropped Sorghum bicolor and Acacia saligna were compared with the agroforestry combination in an alley cropping system in semiarid Northern Kenya. Three methods for the calculation of root production were used: the max-min, balancing-transfer and compartment-flow method. The highest root biomass was found in the topsoil for all cropping systems, though trees had a deeper root system. Trees and crops had a similar amount of below-ground biomass during the vegetation period (0.3 and 0.4 Mg DM ha^-1 120 cm^-1), but in the agroforestry combination root biomass was more than the sum of the sole cropped systems (1.1 Mg DM ha^-1 120 cm^-1). The tree system showed a very static root development with little fluctuation between seasons, whereas root biomasses were very dynamic in the crop and tree + crop systems. Root production was highest in the tree + crop combination with 2.1 Mg DM ha^-1 a^-1, with about 50% less in sole cropped trees and crops. Root N input to soil decreased in the order tree + crop>tree>crop system with 13.5, 11.0 and 3.2 kg N ha^-1 a^-1, and cannot be estimated from total below-ground biomass or carbon turnover, as N is accumulated in senescing roots. Such low N input to soil stresses the need for investigating other processes of nutrient input from roots to soil. Areas of highest N input were identified in the topsoil under the tree row in the tree system. Resource utilisation and C and N input to soil were highest with a combination of annual and perennial crops.     

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.     

Biomass and production of fine and coarse roots during regrowth of a disturbed subtropical humid forest in north-east India

Seasonal variation and depthwise distribution of dry matter in roots of different diameter classes and their annual production were studied using sequential core sampling. The investigations were carried out in three stands of a subtropical humid forest of north-east India representing different stages of regrowth after tree cutting. The mean annual standing crop of fine (<2 mm in diameter) and coarse (2–15 mm diameter) roots increased gradually from 5.4 Mg ha^-1 and 0.7 Mg ha^-1 in 7-yr old regrowth to 9.4 Mg ha^-1 and 2.8 Mg ha^-1 in 16-yr old regrowth, respectively. The contribution of fine roots to the total root mass declined from 88% in 7-yr old regrowth to 77% in both 13 and 16-yr old regrowths, while that of coarse roots increased from 12 to 23%. A major portion of fine roots (59–62%) was present in 0–10 cm soil layer, but the coarse roots were concentrated in 10–20 cm soil depth (38–48%). In all the three stands, biomass of both fine and coarse roots followed a unimodal growth curve by showing a gradual increase from spring/pre-rainy season to autumn/post-rainy season. Biomass to necromass ratio increased from 2.5 in the 7-yr old to 3.2 in the 16-yr old stand. The annual fine root production increased from 5.9 Mg ha^-1 to 7.7 Mg ha^-1 and total root production from 7.6 Mg ha^-1 to 14.7 Mg ha^-1 from 7-yr to 16-yr old regrowth.     

Standing fine root mass and production in four Chinese subtropical forests along a succession and species diversity gradient

Background and aims

The influences of succession and species diversity on fine root production are not well known in forests. This study aimed to investigate: (i) whether fine root biomass and production increased with successional stage and increasing tree species diversity; (ii) how forest type affected seasonal variation and regrowth of fine roots.

Methods

Sequential coring and ingrowth core methods were used to measure fine root production in four Chinese subtropical forests differing in successional stages and species diversity.

Results

Fine root biomass increased from 262 g·m^?2 to 626 g·m^?2 with increasing successional stage and species diversity. A similar trend was also found for fine root production, which increased from 86 to 114 g·m^?2 yr ^?1 for Cunninghamia lanceolata plantation to 211–240 g·m^?2 yr ^?1 for Choerospondias axillaries forest when estimated with sequential coring data. Fine root production calculated using the ingrowth core data ranged from 186 g·m^?2 yr ^?1 for C. lanceolata plantation to 513 g·m^?2 yr ^?1 for Lithocarpus glaber – Cyclobalanopsis glauca forest.

Conclusions

Fine root biomass and production increased along a successional gradient and increasing tree species diversity in subtropical forests. Fine roots in forests with higher species diversity exhibited higher seasonal variation and regrowth rate.     

Fine Root Production across a Primary Successional Ecosystem Chronosequence at Mt. Shasta,California

Estimating changes in belowground biomass and production is essential for understanding fundamental patterns and processes during ecosystem development. We examined patterns of fine root production, aboveground litterfall, and forest floor accumulation during forest primary succession at the Mt. Shasta Mudflows ecosystem chronosequence. Fine root production was measured using the root ingrowth cores method over 1 year, and aboveground litterfall was collected over 2 years. Fine root production increased significantly with ecosystem age, but only the youngest ecosystem was significantly different from all of the older ecosystems. Root production was 44.5 ± 13.3, 168.3 ± 20.6, 190.5 ± 33.8, and 236.3 ± 65.4 g m⁻² y⁻¹ in the 77, 255, 616, and >850-year-old ecosystems, respectively. Generally, aboveground litterfall and forest floor accumulation did not follow the same pattern as root production. The relative contribution of fine root production to total fine detrital production increased significantly with ecosystem age, from 14 to 49%, but only the youngest ecosystem was significantly different from all of the older ecosystems. Fine root production was significantly correlated with some measures of soil fertility but was not correlated with leaf or total litterfall, or forest floor accumulation. It was best predicted by soil N concentration alone, but this relationship may not be causal, as soil N concentration was also correlated with ecosystem age. For the oldest ecosystem, fine root production was also measured using the sequential intact cores/compartment-flow model method, and the difference between the two estimates was not significant. Our study suggests that the relative contribution of fine roots to fine detrital production, and hence to soil organic matter accumulation, may increase during forest primary succession.     

Fine Root Productivity and Turnover in Two Evergreen Central Himalayan Forests

Fine root production and mortality in central Himalayan evergreenforests consisting of Quercus leucotrichophora (banj oak) andPinus roxburghii (chir pine) were measured. Fine root productionand mortality decreased with increasing soil depth. Annual fineroot production was higher in the broadleafed forest than inthe coniferous forest, across months and seasons (1.3 and 1.5-timesmore living and dead root biomass, respectively in banj oakthan in chir pine). Live fine root production was 2508 kg ha^-1year^-1inchir pine forest and 3631 kg ha^-1year^-1in banj oak forest. Deadfine roots accumulated at a rate of 1197 and 1525 kg ha^-1year^-1inchir pine and in banj oak forest, respectively. In both forests,the greatest fine root production was recorded in the rainyseason followed by summer and winter seasons. Both soil androot nitrogen concentration decreased with increasing soil depth.Nitrogen uptake was higher in banj oak forest (12.1 kg ha^-1year^-1)than chir pine forest (7.2 kg ha^-1year^-1).Copyright 1999 Annalsof Botany Company Fine root production, fine roots, necromass, banj oak, chir pine, Quercus leucotrichophora , Pinus roxburghii .     

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.     

Fine root dynamics along a 2,000-m elevation transect in South Ecuadorian mountain rainforests

Fine root turnover plays an important role in the cycling of carbon and nutrients in ecosystems. Not much is known about fine root dynamics in tropical montane rainforests, which are characterized by steep temperature gradients over short distances. We applied the minirhizotron technique in five forest stands along an elevational transect between 1,050 and 3,060 m above sea level in a South Ecuadorian montane rainforest in order to test the influence of climate and soil parameters on fine root turnover. Turnover of roots with diameter <?2.0 mm was significantly higher in the lowermost and the uppermost stand (0.9 cm cm^?1 year^?1) than in the three mid-elevation stands (0.6 cm cm^?1 year^?1). Root turnover of finest roots (d?<?0.5 mm) was higher compared to the root cohort with d?<?2.0 mm, and exceeded 1.0 cm cm^?1 year^?1 at the lower and upper elevations of the transect. We propose that the non linear altitudinal trend of fine root turnover originates from an overlapping of a temperature effect with other environmental gradients (e.g. adverse soil conditions) in the upper part of the transect and that the fast replacement of fine roots is used as an adaptive mechanism by trees to cope with limiting environmental conditions.     

Responses of fine roots and soil N availability to short-term nitrogen fertilization in a broad-leaved Korean pine mixed forest in northeastern China

Knowledge of the responses of soil nitrogen (N) availability, fine root mass, production and turnover rates to atmospheric N deposition is crucial for understanding fine root dynamics and functioning in forest ecosystems. Fine root biomass and necromass, production and turnover rates, and soil nitrate-N and ammonium-N in relation to N fertilization (50 kg N ha(-1) year(-1)) were investigated in a temperate forest over the growing season of 2010, using sequential soil cores and ingrowth cores methods. N fertilization increased soil nitrate-N by 16% (P<0.001) and ammonium-N by 6% (P<0.01) compared to control plots. Fine root biomass and necromass in 0-20 cm soil were 13% (4.61 vs. 5.23 Mg ha(-1), P<0.001) and 34% (1.39 vs. 1.86 Mg ha(-1), P<0.001) less in N fertilization plots than those in control plots. The fine root mass was significantly negatively correlated with soil N availability and nitrate-N contents, especially in 0-10 cm soil layer. Both fine root production and turnover rates increased with N fertilization, indicating a rapid underground carbon cycling in environment with high nitrogen levels. Although high N supply has been widely recognized to promote aboveground growth rates, the present study suggests that high levels of nitrogen supply may reduce the pool size of the underground carbon. Hence, we conclude that high levels of atmospheric N deposition will stimulate the belowground carbon cycling, leading to changes in the carbon balance between aboveground and underground storage. The implications of the present study suggest that carbon model and prediction need to take the effects of nitrogen deposition on underground system into account.     

Comprehensive assessments of root biomass and production in a Kobresia humilis meadow on the Qinghai-Tibetan Plateau

Alpine meadow covers ca. 700,000 km² with an extreme altitude range from 3200 m to 5200 m. It is the most widely distributed vegetation on the vast Qinghai-Tibetan Plateau. Previous studies suggest that meadow ecosystems play the most important role in both uptake and storage of carbon in the plateau. The ecosystem has been considered currently as an active “CO₂ sink”, in which roots may contribute a very important part, because of the large root biomass, for storage and translocation of carbon to soil. To bridge the gap between the potential importance and few experimental data, root systems, root biomass, turnover rate, and net primary production were investigated in a Kobresia humilis meadow on the plateau during the growing season from May to September in 2008 and 2009. We hypothesized that BNPP/NPP of the alpine meadow would be more than 50%, and that small diameter roots sampled in ingrowth cores have a shorter lifespan than the lager diameter roots, moreover we expected that roots in surface soils would turn over more quickly than those in deeper soil layers. The mean root mass in the 0–20 cm soil layer, investigated by the sequential coring method, was 1995?±?479 g?m^?2 and 1595?±?254 g?m^?2 in growing season of 2008 and 2009, respectively. And the mean fine root biomass in ingrowth cores of the same soil layer was 119?±?37 g?m^?2 and 196?±?45 g?m^?2 in the 2 years. Annual total NPP was 12387 kg?ha^?1?year^?1, in which 53% was allocated to roots. In addition, fine roots accounted for 33% of belowground NPP and 18% of the total NPP, respectively. Root turnover rate was 0.52 year^?1 for bulk roots and 0.74 year^?1 for fine roots. Furthermore, roots turnover was faster in surface than in deeper soil layers. The results confirmed the important role of roots in carbon storage and turnover in the alpine meadow ecosystem. It also suggested the necessity of separating fine roots from the whole root system for a better understanding of root turnover rate and its response to environmental factors.     

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

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

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.     

Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.