Competition in tree row agroforestry systems. 2. Distribution, dynamics and uptake of soil inorganic N

The effect of tree row species on the distribution of soil inorganic N and the biomass growth and N uptake of trees and crops was investigated beneath a Grevillea robustaA. Cunn. ex R. Br. (grevillea) tree row and Senna spectabilisDC. (senna) hedgerow grown with Zea mays L. (maize) and a sole maize crop, during one cropping season. The hypothesis was that a tree with a large nutrient uptake would have a greater competitive effect upon coexisting plants than a tree that takes up less and internally cycles nutrients. The field study was conducted on a kaolinitic Oxisol in the sub-humid highlands of western Kenya. Soil nitrate and ammonium were measured to 300 cm depth and 525 cm distance from the tree rows, before and after maize cropping. Ammonium concentrations were small and did not change significantly during the cropping season. There was > 8 mg nitrate kg^–1 in the upper 60 cm and at 90–180 cm depth at the start of the season, except within 300 cm of the senna hedgerow where concentrations were smaller. During the season, nitrate in the grevillea-maize system only decreased in the upper 60 cm, whereas nitrate decreased at almost every depth and distance from the senna hedgerow. Inorganic N (nitrate plus ammonium) decreased by 94 kg ha^–1 in the senna-maize system and 33 kg ha^–1 in the grevillea-maize system.The aboveground N content of the trees increased by 23 kg ha^–1 for grevillea and 39 kg ha^–1 for senna. Nitrogen uptake by maize was 85 kg ha^–1 when grown with grevillea and 65 kg ha^–1 with senna. Assuming a mineralisation input of 50 kg N ha^–1season^–1, the decrease in inorganic soil N approximately equalled plant N uptake in the grevillea-maize system, but exceeded that in the senna-maize system. Pruning and litter fall removed about 14 kg N ha^–1 a^–1 from grevillea, and > 75 kg N ha^–1 a^–1 from senna. The removal of pruned material from an agroforestry system may lead to nutrient mining and a decline in productivity.     

Root distributions partially explain 15N uptake patterns in Gliricidia and Peltophorum hedgerow intercropping systems

The relative distributions of tree and crop roots in agroforestry associations may affect the degree of complementarity which can be achieved in their capture of below ground resources. Trees which root more deeply than crops may intercept leaching nitrogen and thus improve nitrogen use efficiency. This hypothesis was tested by injection of small doses of (¹⁵NH₄)₂SO₄ at 21.8 atom% ¹⁵N at different soil depths within established hedgerow intercropping systems on an Ultisol in Lampung, Indonesia. In the top 10 cm of soil in intercrops of maize and trees, root length density (L_rv) of maize was greater than that of Gliricidia sepium trees, which had greater L_rv in this topsoil layer than Peltophorum dasyrrachis trees. Peltophorum trees had a greater proportion of their roots in deeper soil layers than Gliricidia or maize. These vertical root distributions were related to the pattern of recovery of ¹⁵N placed at different soil depths; more ¹⁵N was recovered by maize and Gliricidia from placements at 5 cm depth than from placements at 45 or 65 cm depth. Peltophorum recovered similar amounts of ¹⁵N from placements at each of these depths, and hence had a deeper N uptake distribution than Gliricidiaor maize. Differences in tree L_rv across the cropping alley were comparatively small, and there was no significant difference (P<0.05) in the uptake of ¹⁵N placed in topsoil at different distances from hedgerows. A greater proportion of the ¹⁵N recovered by maize was found in grain following ¹⁵N placement at 45 cm or 65 cm depth than following placement at 5 cm depth, reflecting the later arrival of maize roots in these deeper soil layers. Thus trees have an important role in preventing N leaching from subsoil during early crop establishment, although they themselves showed a lag phase in ¹⁵N uptake after pruning. Residual ¹⁵N enrichment in soil was strongly related to application depth even 406 days after ¹⁵N placement, demonstrating the validity of this approach to mapping root activity distributions.     

Alley cropping with Senna siamea in South-western Nigeria: II. Dry matter,total N,and urea-derived N dynamics of the Senna and maize roots

Crop and tree roots are crucial in the nutrient recycling hypotheses related to alley cropping systems. At the same time, they are the least understood components of these systems. The biomass, total N content and urea-derived N content of the Senna and maize roots in a Senna-maize alley cropping system were followed for a period of 1.5 years (1 maize-cowpea rotation followed by 1 maize season) to a depth of 90 cm, after the application of ¹⁵N labeled urea. The highest maize root biomass was found in the 0–10 cm layer and this biomass peaked at 38 and 67 days after planting the 1994 maize (DAP) between the maize rows (112 kg ha^–1, on average) and at 38, 67 and 107 DAP under the maize plants (4101 kg ha^–1, on average). Almost no maize roots were found below 60 cm at any sampling date. Senna root biomass decreased with time in all soil layers (from 512 to 68 kg ha^–1 for the 0–10 cm layer between 0 and 480 DAP). Below 10 cm, at least 62% of the total root biomass consisted of Senna roots and this value increased to 87% between 60 and 90 cm. Although these observations support the existence of a Senna root `safety net' between the alleys which could reduce nutrient leaching losses, the depth of such a net may be limited as the root biomass of the Senna trees in the 60–90 cm layer was below 100 kg ha^–1, equivalent to a root length density of only < 0.05 cm cm^–3. The proportion of maize root N derived from the applied urea (%Ndfu) decreased significantly with time (from 21% at 21 DAP to 8% at 107 DAP), while %Ndfu of the maize roots at the second harvest (480 DAP) was only 0.6%. The %Ndfu of the Senna roots never exceeded 4% at any depth or sampling time, but decreased less rapidly compared to the %Ndfu of the maize roots. The higher %Ndfu of the maize roots indicates that maize is more efficient in retrieving urea-derived N. The differences in dynamics of the %Ndfu also indicate that the turnover of N through the maize roots is much faster than the turnover of N through the Senna roots. The recovery of applied urea-N by the maize roots was highest in the top 0–10 cm of soil and never exceeded 0.4% (at 38 DAP) between the rows and 7.1% (at 67 DAP) under the rows. Total urea N recovery by the maize roots increased from 1.8 to 3.2% during the 1994 maize season, while the Senna roots never recovered more than 0.8% of the applied urea-N at any time during the experimental period. These values are low and signify that the roots of both plants will only marginally affect the total recovery of the applied urea-N. Measurement of the dynamics of the biomass and N content of the maize and Senna roots helps to explain the observed recovery of applied urea-N in the aboveground compartments of the alley cropping system.     

Sieve size effects on root length and biomass measurements of maize (Zea mays) and Grevillea robusta

The purpose of this study was to investigate the effects of different mesh sizes on the recovery of root length and biomass and to determine whether the degree of recovery was influenced by plant species and sample location. Sieves of 2.0, 1.0, 0.5 and 0.25 mm (4.0, 1.0, 0.25 and 0.06 mm2) mesh sizes were used to recover and measure the root length and biomass of Zea mays L. (maize) at 0–15 cm and 30–45 cm depths and of Grevillea robusta A. Cunn. ex R. Br. (grevillea) at the same depths 1.0 m and 4.5 m from a line of grevillea trees. At 0–15 cm, the coarser sieves (sum collected with 2.0 and 1.0 mm sieves) recovered approximately 80% of the total root biomass measured, but only 60% of the root length. The proportion of total maize root length and biomass recovered by the coarser sieves decreased with soil depth. The proportion of total grevillea root length recovered by the coarser sieves was similar at the two soil depths, but increased slightly with distance from the tree line. The ≥ 0.5 mm sieves recovered between 93 and 96% of grevillea and maize root biomass and between 73 and 98% of their root length, depending on the sample location. Roots passing through the 0.5 mm sieve, but recovered by the 0.25 mm sieve were about 20% of total maize root length and grevillea root length at 1.0 m from the tree line but < 5% of the total grevillea root length at 4.5 m from the tree. Roots passing through the 0.5 mm sieve but recovered by the 0.25 mm sieve contributed only slightly to root biomass. Although the ≥ 0.5 mm sieves provided adequate measurements of root biomass, the ≥ 0.25 mm sieves were required for accurate measurement of fine root length. There was no universal correction for root length and biomass underestimation when large sieve sizes were used because the proportions of length and biomass recovered depended on the plant species and on soil depth and distance from the plant. This revised version was published online in August 2006 with corrections to the Cover Date.     

Roots,soil water and crop yield: tree crop interactions in a semi-arid agroforestry system in Kenya

Variations in soil water, crop yield and fine roots of 3–4 year-old Grevillea robusta Cunn. and Gliricidia sepium (Jacq.) Walp. growing in association with maize (Zea mays L.) were examined in semiarid Kenya during the long rains of 1996 and 1997. Even although tree roots penetrated more deeply than maize roots, maximum root length densities for both tree species and maize occurred in the top 200 mm of the soil profile where soil moisture was frequently recharged by rains. Populations of roots in plots containing trees were dominated by tree roots at the beginning of the growing season but because tree roots died and maize root length increased during the cropping season, amounts of tree and maize roots were similar at the end of the season. Thus, there was evidence of temporal separation of root activity between species, but there was no spatial separation of the rooting zones of the trees and crops within that part of the soil profile occupied by crop roots. Tree root length density declined with increasing distances from rows of trees and with depth in the soil profile. Although Grevillea trees were largest, plots containing G. sepium trees always contained more tree roots than plots containing G. robusta trees and Gliricidia was more competitive with maize than Grevillea. Overall, Gliricidia reduced crop yield by about 50% and Grevillea by about 40% relative to crop yield in control plots lacking trees and reductions of crop yield were greatest close to trees. There was less soil moisture in plots containing trees than in control plots. Such difference between control plots and plots containing trees were maximal at the end of the dry season and there was always less soil moisture close to trees than elsewhere in the plots. Plots containing Gliricidia trees contained less soil water than plots containing Grevillea trees.     

Effects of herbicide chlorsulfuron on growth and nutrient uptake parameters of wheat genotypes differing in Zn-efficiency

Prunings of Calliandra calothyrsus, Grevillea robusta, Leucaena diversifolia and farm yard manure were applied each cropping season at 3 and 6 t dry matter ha⁻¹ to an Oxisol in Burundi. The field plots also received basal applications of nitrogen (N), phosphorus (P) and potassium (K). Application of the tree prunings or farm yard manure decreased the concentration of monomeric inorganic aluminium (Al) in soil solution from 2.92 mg Al dm⁻³ in the control plots to 0.75 mg Al dm⁻³ in the plots receiving 6 t ha⁻¹ Calliandra prunings. The other organic materials also decreased the concentration of monomeric inorganic aluminium in the soil solution. The lowered Al concentration led to a corresponding decrease in the percentage Al saturation of the 0–10 cm soil layer from 80% to 68%. Grain yields of maize and beans were strongly inversely related to the percentage Al saturation of the soil. This confirms that soil acidity was the main constraint to maize and beans production. The yield improvement was mainly attributed to the ameliorating effects of the organic matter application on Al toxicity. The nutrient content had less effect presumably because of fertilizer use. In the best treatments, the yield of maize increased from 0.9 to 2.2 t ha⁻¹ and the corresponding beans yield increased from 0.2 to 1.2 t ha⁻¹. A C Borstlap Section editor     

Distribution,length and weight of roots in young plantations ofEucalyptus grandis W. Hill ex Maiden irrigated with recycled water

Summary The root systems ofEucalyptus grandis W. Hill ex Maiden, irrigated with recycled municipal effluent at two sites in north-western Victoria, Australia, were studied by excavation and coring. Trees at Robinvale were four years-old and were irrigated using micro-sprays that covered only 70% of the ground surface area, whereas at Mildura, effuent was uniformly was uniformly applied to six years-old trees by flood and sprinkler irrigation. At Mildura where roots were excavated from a 2.80×2.80×1.20 m block of soil, a total root length of 1193 m.m⁻² and a total root weight of 3.1 kg m⁻² were estimated in the top metre. For roots >1 mm diameter, 77% of intercepts were at 0–30 cm, whereas only 50% were in the 50–100 cm soil horizon. At both sites where roots in the top 30 cm were studied by coring, the vertical distributions of root intercepts, length and weight were similar. Root length was greatest in the 0–10 cm soil horizon at both sites, and intercepts of roots <1 mm diameter comprised 73% and 81% of all roots at Mildura and Robinvale respectively. Roots <1 mm diameter contributed 85% of total length at both sites, but only 19% and 21% of total weight at Mildura and Robinvale respectively. The horizontal distribution of roots differed at the two sites. With uniform application of effuent at Mildura, root intercepts and length were concentrated in the centre of the irrigation bay, but at Robinvale, the concentration occurred closer to the tree row due mainly to the different method of irrigation. Root weight at both sites was highest within 50 cm of the tree row. Root densities of 0.11 to 0.57 cm cm⁻³ were estimated in the two plantations; these were similar to root densities measured inPinus radiata D. Don plantations up to 46 months old, but were considerably lower than those estimated for pastures. The implications of the results for the management of irrigated plantations of eucalypts are discussed.     

Inoculum size and auxin concentration influence the growth of adventitious roots and accumulation of ginsenosides in suspension cultures of ginseng ( Panax ginseng C.A. Meyer)

The effect of the root-inoculum size and axuin concentration on growth of adventitious roots and accumulation of ginsenosides were studied during suspension cultures of ginseng (Panax ginseng C.A. Meyer). Of the various concentrations of indole-3-butyric acid (IBA) and γ-naphthaleneacetic acid (NAA) used as supplementary growth regulators along with Murashige and Skoog medium, 25 μM IBA was found suitable for lateral root induction and growth, as well as accumulation of ginsenosides. Inoculum size of 5 g L⁻¹ was found suitable for optimal biomass (10.5 g L⁻¹ dry biomass) and ginsenosides (5.4 mg g⁻¹ DW) accumulation. Of the various length of root inocula tested (chopped to 1–3, 4–6, 7–10 mm and un-chopped), root inocula of 7–10 mm was found suitable for biomass and ginsenoside accumulation.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

Effects of Temperature and Body Size on the Swimming Speed of Larval and Juvenile Atlantic Cod (Gadus Morhua): Implications for Individual-based Modelling

Synopsis The routine swimming speed (S) of three groups of 4, 9 and 32 cm total length (L_T) juvenile cod (Gadus morhua) was quantified in the laboratory at 6 – 10 different temperatures (T) between 3.2 and 16.7°C. At temperatures between 5 and 15°C, mean group S increased exponentially with increasing T (S=a e^bT) and the effect of temperature (b = 0.082, Q₁₀ = 2.27) was not significantly different among the groups (over the 8-fold difference in fish sizes of early- and post-settlement juveniles). Differences in mean S among individuals within each group were quite large (coefficient of variation = 40 – 80%). Swimming data for juveniles and those collected for groups of 0.4, 0.7 and 0.9 cm standard length (L_S) larvae were combined to assess the effect of body size on S. At 8°C, S (mm s⁻¹) increased with L_S (mm) according to: S = 0.26L_S^Φ−5.28L_S⁻¹, where Φ = 1.55L_S^−0.08. Relative S (body lengths s⁻¹) was related to L_S by a dome-shaped relationship having a maximum value (0.49 body lengths s⁻¹) at 18.5 – 19 mm L_S corresponding to the sizes of fish at the end of larval-juvenile metamorphosis. Previous larval cod IBM’s using a cruise-predator mode likely overestimated rates of foraging (prey searching and encounters) by a factor of ~2, whereas foraging rates in pause-travel models are closer to estimates of swimming velocities obtained in this and other laboratory studies.     

Soil O2 and CO2 effects on root respiration of cacti

Through use of a recently developed technique that can measure CO₂ exchange by individual attached roots, the influences of soil O₂ and CO₂ concentrations on root respiration were determined for two species of shallow-rooted cacti that typically occur in porous, well-drained soils. Although soil O₂ concentrations in the rooting zone in the field were indistinguishable from that in the ambient air (21% by volume), the CO₂ concentrations 10 cm below the soil surface averaged 540 μLL⁻¹ for the barrel cactusFerocactus acanthodes under dry conditions and 2400 μLL⁻¹ under wet conditions in a loamy sand. For the widely cultivated platyopuntiaOpuntia ficus-indica in a sandy clay loam, the CO₂ concentration at 10 cm averaged 1080 μLL⁻¹ under dry conditions and 4170 μLL⁻¹ under wet conditions. For both species, the respiration rate in the laboratory was zero at 0% O₂ and increased to its maximum value at 5% O₂ for rain roots (roots induced by watering) and 16% O₂ for established roots. Established roots ofO. ficus-indica were slightly more tolerant of elevated CO₂ than were those ofF. acanthodes, 5000 μLL⁻¹ inhibiting respiration by 35% and 46%, respectively. For both species, root respiration was reduced to zero at 20,000 μLL⁻¹ (2%) CO₂. In contrast to the reversible effects of 0% O₂, inhibition by 2% CO₂ was irreversible and led to the death of cortical cells in established roots in 6 h. Although the restriction of various cacti and other CAM plants to porous soils has generally been attributed to their requirement for high O₂ concentrations, the present results indicate that susceptibility of root respiration to elevated soil CO₂ concentrations may be more important.     

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]     

Dinitrogen-fixation by three neotropical agroforestry tree species under semi-controlled field conditions

Cultivating dinitrogen-fixing legume trees with crops in agroforestry is a relatively common N management practice in the Neotropics. The objective of this study was to assess the N₂ fixation potential of three important Neotropical agroforestry tree species, Erythrina poeppigiana, Erythrina fusca, and Inga edulis, under semi-controlled field conditions. The study was conducted in the humid tropical climate of the Caribbean coastal plain of Costa Rica. In 2002, seedlings of I. edulis and Vochysia guatemalensis were planted in one-meter-deep open-ended plastic cylinders buried in soil within hedgerows of the same species. Overall tree spacing was 1 × 4 m to simulate a typical alley-cropping design. The ¹⁵N was applied as (NH₄)₂SO₄ at 10% ¹⁵N atom excess 15 days after planting at the rate of 20 kg [N] ha⁻¹. In 2003, seedlings of E. poeppigiana, E. fusca, and V. guatemalensis were planted in the same field using the existing cylinders. The ¹⁵N application was repeated at the rate of 20 kg [N] ha⁻¹ 15 days after planting and 10 kg [N] ha⁻¹ was added three months after planting. Trees were harvested 9 months after planting in both years. The ¹⁵N content of leaves, branches, stems, and roots was determined by mass spectrometry. The percentage of atmospheric N fixed out of total N (%N_f) was calculated based on ¹⁵N atom excess in leaves or total biomass. The difference between the two calculation methods was insignificant for all species. Sixty percent of I. edulis trees fixed N₂; %N_f was 57% for the N₂-fixing trees. Biomass production and N yield were similar in N₂-fixing and non-N₂-fixing I. edulis. No obvious cause was found for why not all I. edulis trees fixed N₂. All E. poeppigiana and E. fusca trees fixed N₂; %N_f was ca. 59% and 64%, respectively. These data were extrapolated to typical agroforestry systems using published data on N recycling by the studied species. Inga edulis may recycle ca. 100 kg ha⁻¹ a⁻¹ of N fixed from atmosphere to soil if only 60% of trees fix N₂, E. poeppigiana 60–160 kg ha⁻¹ a⁻¹, and E. fusca ca. 80 kg ha⁻¹ a⁻¹.     

Fine Root Distribution in Dehesas of Central-Western Spain

A Dehesa is a structurally complex agro-silvo-pastoral system where at least two strata of vegetation, trees and herbaceous plants coexist. We studied the root distribution of trees (Quercus ilex L.) and herbaceous plants, in order to evaluate tree and crops competition and complementarity in Dehesas of Central Western Spain. 72 soil cores of 10 cm diameter (one to two metre deep) were taken out around 13 trees. Seven trees were intercropped with Avena sativa L. and six trees were in a grazed pasture dominated by native grasses. Soil coring was performed at four distances from the tree trunks, from 2.5 (beneath canopy) till 20 m (out of the canopy). Root length density (RLD) of herbaceous plants and trees was measured using the soil core-break method. Additionally, we mapped tree roots in 51 profiles of 7 recently opened road cuts, located between 4 and 26 m of distance from the nearest tree. The depth of the road cuts varied between 2.5 and 5.5 m. Herbaceous plant roots were located mostly in the upper 30 cm, above a clayey, dense soil layer. RLD of herbaceous plants decreased exponentially with depth until 100 cm depth. Holm-oak showed a much lower RLD than herbs (on average, 2.4 versus 23.7 km m⁻³, respectively, in the first 10 cm of the soil depth). Tree RLD was surprisingly almost uniform with depth and distance to trees. We estimated a 5.2 m maximum depth and a 33 m maximum horizontal extension for tree roots. The huge surface of soil explored by tree roots (even 7 times the projection of the canopy) could allow trees to meet their water needs during the dry Mediterranean summers. The limited vertical overlap of the two root profiles suggests that competition for soil resources between trees and the herbaceous understorey in the Dehesa is probably not as strong as usually assumed.     

Effects of aluminium and mineral nutrition on growth and chemical composition of hydroponically grown seedlings of five different forest tree species

Forest die-back and impaired tree vitality have frequently been ascribed to Al-toxicity and Al-induced nutritional disorders due to increased acidification of forest soils. Therefore, in this experiment effects of Al were studied on growth and nutrient uptake with seedlings of five different forest tree species. During growth in culture solutions with and without Al all five species proved to be very Al-tolerant, despite high accumulation of Al in roots. In the coniferous evergreens Douglas-fir and Scots pine shoot as well as root Al concentrations were significantly higher than in the deciduous broad-leaved species oak and birch. Larch showed intermediate Al levels. In none of the five species did Al reduce nutrient concentrations or the Ca/Al ratio to values below the critical level. Besides differences in Al accumulation, coniferous and broad-leaved species also differed with respect to uptake and assimilation of nitrogen. Due to extra NH ₄ ⁺ uptake, oak and birch showed a much higher N uptake and higher NH ₄ ⁺ preference than the coniferous species. Especially with oak this high NH ₄ ⁺ preference in combination with a low specific root surface area resulted in a high root proton efflux density. In comparison to both broad-leaved trees and Scots pine the NO ₃ ⁻ reduction capacity of larch and Douglas-fir was extremely low. This may have important consequences for both species if grown in NO ₃ ⁻ -rich soils.     

Dependence of the aboveground CO2 exchange rate on tree size in field-grown hinoki cypress (Chamaecyparis obtusa)

The CO₂ exchange of the aboveground parts for five different-sized 17-year-old (as of 1991) hinoki cypress (Chamaecyparis obtusa) trees growing in the field was non-destructively measured over one year, using an open CO₂ exchange system. The CO₂ exchange of individual trees decreased with decreasing tree sizes, such as aboveground phytomass, leaf mass and leaf area. However, the CO₂ exchange abruptly decreased near the smallest-suppressed sample tree. The size dependence was well described by a generalized power function. The annual gross photosynthesis of individual trees was proportional to the square root of leaf mass or leaf area. The dependence of CO₂ exchange on annual phytomass increment was described by a simple power function with an exponent value less than unity, suggesting that CO₂ exchange per unit of phytomass increment was lower in larger-sized trees than in smaller-sized trees. The mean photosynthetic activity of a tree, i.e., gross photosynthesis per unit of leaf area, slightly increased to its highest value with decreasing leaf area and then decreased abruptly near the smallest sample tree. The maximum value of mean photosynthetic activity was estimated to be 2.85 kg CO₂ m⁻² year⁻¹ for a leaf area of 1.56 m² tree⁻¹. The ratio of mean photosynthetic activity to the maximum photosynthetic activity was the highest in an intermediate tree and decreased gradually toward larger-sized trees by ca. 60% and also decreased toward the smallest suppressed tree by ca. 35%.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

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.     

A Method to Estimate Practical Radial Oxygen Loss of Wetland Plant Roots

The estimation of practical radial oxygen loss (ROL) of wetland plant roots was attempted in this study. We have devised a new method to measure ROL of wetland plant roots. The whole root system was bathed in an anoxic nutrient solution. Oxygen released from the root was removed immediately by introducing oxygen-free nitrogen gas (O₂ < 4 nmol L⁻¹) to mimic natural habitats where released oxygen is consumed rapidly due to chemical and biological oxidation processes. Oxygen removed from the root-bathing chamber was simultaneously detected colorimetrically by use of the highly oxygen-sensitive anthraquinone radical anion (AQ·⁻) in a cell outside the root-bathing chamber, which decolorized by a rapid reaction with oxygen. An emergent macrophyte Typha latifolia L. was incubated, and its ROL was measured by both the new method and one of the conventional methods, the closed chamber/electrode method, by which the ROL of Typha latifolia L. had not yet been measured. The new method succeeded in detecting the ROL, whereas the conventional method was not able to detect oxygen, due to the level being below the detection limit of the oxygen electrode. The oxygen supply via the seedlings of Typha latifolia L. was ca. 10 times higher compared with control measurements without plant. Light illumination significantly enhanced the ROL of Typha latifolia L. (0.33 nmol O₂ g⁻¹ root dry weight s⁻¹ under light and 0.18 nmol O₂ g⁻¹ root dry weight s⁻¹ in the dark). Theses values fall between those previously reported by the closed chamber/titanium citrate method and the open chamber/electrode method.     

Influence of agitation and aeration on growth and antibiotic production by <Emphasis Type="Italic">Xenorhabdus nematophila</Emphasis>

The effect of agitation and aeration on the growth and antibiotic production by Xenorhabdus nematophila YL001 grown in batch cultures were investigated. Efficiency of aeration and agitation was evaluated through the oxygen mass transfer coefficient (K _L a). With increase in K _L a, the biomass and antibiotic activity increased. Activity units of antibiotic and dry cell weight were increased to 232 U ml⁻¹ and 19.58 g l⁻¹, respectively, productivity in cell and antibiotic was up more than 30% when K _L a increased from 115.9 h⁻¹ to 185.7 h⁻¹. During the exponential growth phase, DO concentration was zero, the oxygen supply was not sufficient. So, based on process analysis, a three-stage oxygen supply control strategy was used to improved the DO concentration above 30% by controlling the agitation speed and aeration rate. The dry cell weight and activity units of antibiotic were further increased to 24.22 g l⁻¹ and 249 U ml⁻¹, and were improved by 24.0% and 7.0%, compared with fermentation at a constant agitation speed and a constant aeration rate (300 rev min⁻¹, 2.5 l min⁻¹).