Green manure production of Azolla microphylla and Sesbania rostrata and their long-term effects on rice yields and soil fertility

Summary Azolla spp. and Sesbania spp. can be used as green manure crops for wetland rice. A long-term experiment was started in 1985 to determine the effects of organic and urea fertilizers on wetland rice yields and soil fertility. Results of 10 rice croppings are reported. Azolla sp. was grown for 1 month and then incorporated before transplanting the rice and 3–4 weeks after transplanting the rice. Sesbania rostrata was grown for 7–9 weeks and incorporated only before transplanting the rice. Sesbania sp. grew more poorly before dry season rice than before wet season rice. Aeschynomene afraspera, which was used in one dry season rice trial, produced a larger biomass than the Sesbania sp. The quantity of N produced by the Azolla sp. ranged from 70 to 110 kg N ha^-1. The Sesbania sp. produced 55–90 kg N ha^-1 in 46–62 days. Rice grain yield increases in response to the green manure were 1.8–3.9 t ha^-1, similar to or higher than that obtained in response to the application of 60 kg N ha^-1 as urea. Grain production per unit weight of absorbed N was lower in the green manure treatments than in the urea treatment. Without N fertilizer, N uptake by rice decreased as the number of rice crops increased. For similar N recoveries, Sesbania sp. required a lower N concentration than the Azolla sp. did. Continuous application of the green manure increased the organic N content in soil on a dry weight basis, but not on a area basis, because the application of green manure decreased soil bulk density. Residual effects in the grain yield and N uptake of rice after nine rice crops were found with a continuous application of green manure but not urea.     

Effect of phosphorus fertilization on blue-green algal inoculum production and nitrogen yield under field conditions

Summary Field experiments were conducted to assess the effects of the application of P on growth and N yield of inoculated and indigenous blue-green algae (BGA). Addition of 17.4 kg P ha^–1 in split applications led to the highest BGA biomass and N yield, 162 kg dry weight ha^–1 and 6 kg N ha^–1 per 15 days, respectively. When inoculum of Aulosira spp., Aphanothece spp., Gloeotrichia spp. were compared separately, Gloeotrichia spp. grew faster but Aulosira spp. fixed more N. The growth rate and N yield of Aulosira spp. decreased with high P applications, although growth continued until the application of 34.8 kg P ha^–1. The effects of P on inoculum production by local species compared with those collected from other states showed the superiority of the local culture. Applications of P also enhanced the growth and N yield of indigenous BGA, with Wollea spp. showing the best results.     

Effects of crop growth and soil treatments on microbial C,N, and P in dry tropical arable land

We studied the dynamics of microbial C, N, and P in soil cropped with rice (Oryza sativa) and lentils (Lens culinaris) in a dryland farming system. The crop biomass and grain yield were also studied. The microbial biomass and its N and P contents were larger under the lentil than under the rice crop. Microbial nutrients decreased as the crops grew and then increased again. Farmyard manure and NPK fertilizer applications increased the level of microbial nutrients, crop biomass, and grain yield by 35–80%, 55–85%, and 74–86%, respectively. However, these applications had no significant effect on most of the soil physicochemical properties in the short term. The microbial biomass was correlated with the crop biomass and grain yield. The calculated flux of N and P through the microbial biomass ranged from 30–45 and 10–19 kg ha^-1 year^-1, respectively. Cultivation of a cereal crop followed by a leguminous crop sustains higher levels of microbial nutrients and hence greater fertility in impoverished tropical arable soils. The soil microbial biomass appears to contribute significantly to crop productivity by releasing nutrients, and applications of manure, either alone or with fertilizers, promote this effect more strongly than the application of NPK fertilizers alone.     

Sensitivity of forest-floor mosses in boreal forests to nitrogen and sulphur deposition

The response of forest-floor mosses to deposition of nitrogen (N) and sulphur (S) was examined in field conditions in a 60-year-old Norway spruce (Picea abies Karst.) stand in southern Finland. The experimental plots received nitrogen (25 kg N ha^–1) and sulphur (30 kg S ha^–1) as ammonium sulphate once a year for 4 years.The dominant moss species on the site were Pleurozium schreberi (Mitt.) and Dicranum polysetum (Sw.). The biomass of the dominant moss species was decreased significantly by N and S deposition during the study period. Due to the addition of N and S, the biomass of Pleurozium schreberi was decreased by 60% and the biomass of Dicranum polysetum by 78%.     

Selection of efficient vesicular-arbuscular mycorrhizal fungi for wetland rice (Oryza sativa L.)

Summary We tested the response of the wetland rice cultivar Prakash to inoculation with ten vescular-arbucular mycorrhizal (VAM) fungi (three selected from the first screening and seven isolated from local paddy fields) in a pot experiment under flooded conditions in order to select the most efficient mycorrhizal fungi to inoculate the rice nursery. A sandy clay loam soil was used as the substrate, fertilized with the recommended N and K levels (100 kg N ha^–1 as ammonium sulphate and 50 kg K ha^–1 as muriate of potash) and half the recommended level of P (25 kg ha^–1 as super phosphate). The inoculation was made into dry nursery beds and the beds were flooded when the seedlings were about 25 cm high, in 15 days. Twenty-eight-day old seedlings were transferred to pots filled with well puddled soil flooded with 5 cm of standing water. Based on the increase in grain yield and total biomass, Glomus intraradices and Acaulospora sp. were considered efficient and suitable for inoculation into rice nurseries.     

Nitrogen contribution to wetland rice by green manuring with Sesbania spp. in an alkaline soil

Summary Two annual species of Sesbania, S. aculeata and Sesbania sp. PL Se-17, were field evaluated as green manure for wetland rice in an alkaline soil. The two species were raised as a catch crop during summer in a wheat-rice rotation, and added as 24.7 and 20.8 t ha^–1 of green matter, 116 and 98 kg N ha^–1, respectively, after 45 days of growth. For the optimum green manuring effect on rice grain yield and N uptake, S. aculeata required 5 days of decomposition (after turning in and before rice transplantation), whereas no decomposition period was necessary for Sesbania sp. PL Se-17. The effect on grain yield and N uptake of rice was equivalent to an application of 122 and 78 kg ha^–1 of chemical N, for the two species, respectively. There was no residual effect of the green manuring on the soil N status after rice harvest.     

Effect of bamboo harvest on dynamics of nutrient pools,N mineralization,and microbial biomass in soil

The effect of harvesting bamboo savanna on the dynamics of soil nutrient pools, N mineralization, and microbial biomass was examined. In the unharvested bamboo site NO _inf3 ^sup- -N in soil ranged from 0.37 to 3.11 mg kg^-1 soil and in the harvested site from 0.43 to 3.67 mg kg^-1. NaHCO₃-extractable inorganic P ranged from 0.55 to 3.58 mg kg^-1 in the unharvested site and from 1.01 to 4.22 mg kg^-1 in the harvested site. Over two annual cycles, the N mineralization range in the unharvested and harvested sites was 0–19.28 and 0–24.0 mg kg^-1 soil month^-1, respectively. The microbial C, N, and P ranges were 278–587, 28–64, and 12–26 mg kg^-1 soil, respectively, with the harvested site exhibiting higher values. Bamboo harvesting depleted soil organic C by 13% and total N by 20%. Harvesting increased N mineralization, resulting in 10 kg ha^-1 additional mineral N in the first 1st year and 5 kg ha^-1 in the 2nd year following the harvest. Microbial biomass C, N and P increased respectively by 10, 18, and 5% as a result of bamboo harvesting.     

Soil water conservation and yield of winter sorghum (Sorghum bicolor L. Moench) as influenced by tillage, organic materials and nitrogen fertilizer in semi-arid tropical India

Soil water and nutrients play an important role in increasing sorghum (Sorghum bicolor L. Moench) yields in the Vertisols of semi-arid tropics during post-rainy season. The effects of tillage practices, organic materials and nitrogen fertilizer on soil properties, water conservation and yield of sorghum were evaluated during winter seasons of 1994–1995 and 1995–1996 on deep Vertisols at Bijapur in the semi-arid tropics of Karnataka State (Zone 3) of south India. Conservation and availability of water and nutrients during different stages of crop growth were increased by deeper tillage resulting in increased grain yield of winter sorghum. Medium and deep tillage increased the grain yield by 23% (1509 kg ha⁻¹) and 57% (1919 kg ha⁻¹) during 1994–1995 and 14% (1562 kg ha⁻¹) and 34% (1835 kg ha⁻¹) during 1995–1996, respectively, over shallow tillage. Water use efficiency increased from shallow (4.90 kg ha⁻¹ mm⁻¹) to deep tillage (7.30 kg ha⁻¹ mm⁻¹). Greater water use efficiency during 1994–1995 as compared to 1995–1996 was attributed to lower consumptive use of water during 1994–1995. Among organic materials, application of Leucaena loppings conserved larger amounts of water and increased winter sorghum yield and water use efficiency. Application of Leucaena loppings increased the winter sorghum grain yield by 9% (mean of 1994–1995 and 1995–1996) as compared to vermicompost. Significantly (P < 0.05) higher water use efficiency of 6.32 kg ha⁻¹ mm⁻¹ was observed in Leucaena loppings incorporated plots compared to 5.72 kg ha⁻¹ mm⁻¹ from vermicompost. Grain yield increased by 245 kg ha⁻¹ with application of 25 kg N ha⁻¹ in 1994–1995, and a further increase in N application to 50 kg ha⁻¹ increased the grain yield by about 349 kg ha⁻¹ in 1995–1996. Deep tillage with application of 25 kg N ha⁻¹ resulted in significantly higher sorghum yield (2047 kg ha⁻¹) than control during 1994–1995. Deep tillage with integrated nutrient management (organic and inorganic N sources) conserved higher amount of soil water and resulted in increased sorghum yields especially during drought years.     

Symbiotic efficiency of Sesbania rostrata and S. cannabina as affected by agronomic practices

A field experiment was conducted to study the N₂ fixation efficiency of Sesbania rostrata and S. cannabina as affected by agronomic practices in semi-arid subtropical climate, Sowing seeds resulted in smaller numbers of nodules, lower dry weight, lower total biomass, less N uptake, and less N₂ fixation for S. rostrata than S. cannabina, while cut-stem planting improved the symbiotic efficiency. Flooding the soil increased the relative humidity of the crop micro-environment by 4–11% and induced early appearance of stem nodules in S. rostrata. Only 67 kg N ha^-1 was fixed by S. rostrata compared to 160 kg N ha^-1 by S. cannabina when normal agronomic practices (sowing and non-flooding) were followed. In contrast, planting stem cuttings and flooding resulted in greater biological N₂ fixation, 307 and 209 kg N ha^-1 by S. rostrata and S. cannabina, respectively. Therefore, S. rostrata can be successfully exploited as a green manure when stem cuttings are planted under flooded conditions.     

Grain yield response of Zea mays (hybrid AE 703) to Azotobacter chroococcum H23

Summary Field experiments were conducted during 1985 and 1986 to examine the effect of Azotobacter chroococcum on the grain yield of maize. Application of 40 kg N ha^–1 plus A. chroococcum caused a significant increase in maize yield. Azotobacter inoculation was more efficient at lower doses (40 kg N ha^–1) than at high doses (80 kg N ha^–1) of urea.     

Effects of fertilizer application and fire regime on soil microbial biomass carbon and nitrogen,and nitrogen mineralization in an Australian subalpine eucalypt forest

The effects of a range of fertilizer applications and of repeated low-intensity prescribed fires on microbial biomass C and N, and in situ N mineralization were studied in an acid soil under subalpine Eucalyptus pauciflora forest near Canberra, Australia. Fertilizer treatments (N, P, N+P, line + P, sucrose + P), and P in particular, tended to lower biomass N. The fertilizer effects were greatest in spring and smaller in summer and late actumn. Low-intensity prescribed fire lowered biomass N at a soil depth of 0–5 cm with the effect being greater in the most frequently burnt soils. No interactions between fire treatments, season, and depth were significant. Only the lime + P and N+P treatments significantly affected soil microbial biomass C contents. The N+P treatment increased biomass C only at 0–2.5 cm in depth, but the soil depth of entire 0–10 cm had much higher (>doubled) biomass C values in the line + P treatment. Frequent (two or three times a year) burning reduced microbial boomass C, but the reverse was true in soils under forest burn at intervals of 7 years. Soil N mineralization was increased by the addition of N and P (alone or in combination), line + P, and sucrose + P to the soil. The same was true for the ratio of N mineralization to biomass N. Soil N mineralization was retarded by repeated fire treatments, especially the more frequent fire treatment where rates were only about half those measured in unburnt soils. There was no relationship between microbial biomass N (kg N ha^-1) and the field rates of soil N mineralization (kg N ha^-1 month^-1). The results suggest that although soil microbial biomass N represents a distinct pool of N, it is not a useful measure of N turnover.     

Effect of soil treatment with cereal straw and method of crop establishment on field pea (Pisum sativum L.) N2 fixation

The effects of soil incorporation with cereal straw (nil, 2.5, 5 and 10 t straw ha^–1) and direct drilling on the proportion and amount of pea N derived from biological N fixation were investigated in three field experiments. Fixed N was determined by¹⁵N dilution using barley as a reference plant. The three sites were on acidic, red clay-loams in the cropping zone of southeastern Australia. Seasonal plant available soil N, as determined by the N accumulated in barley, was 31, 56 and 158 kg N ha^–1, for the three sites. Incorporated straw reduced soil nitrate at sowing by 10–50 kg N ha^–1 (0–30 cm), and 5 or 10 t straw ha^–1 reduced barley uptake of N by 10–38 kg N ha^–1. However, reducing plant available soil N was generally ineffective for increasing the N fixed by pea. Fixed N increased only at the site with the least plant-available N, and only one-third of the increase could be attributed to lower soil N uptake by pea. There was no evidence that direct drilling pea increased fixed N by decreasing crop uptake of soil N. It is proposed that a lower requirement for soil N by pea as compared to barley, and availability of mineral N beneath the soil layer treated with straw, minimise the effectiveness of straw incorporation for increasing the N fixed by pea.     

Phosphorus release during decomposition of crop residues under conventional and zero tillage

Field experiments were conducted at Fort Vermilion (58°23′N 116°02′W), Alberta, to determine phosphorus (P) release patterns from red clover (Trifolium pratense) green manure (GM), field pea (Pisum sativum), canola (Brassica rapa) and monoculture wheat (Triticum aestivum) residues in the 7th and 8th years of conventional and zero tillage. Phosphorus contained in crop residues ranged from 1.5 kg ha⁻¹ in pea to 9.2 kg ha⁻¹ in clover GM, both under zero tillage. The patterns of P release over a 52-week period sometimes varied with tillage, i.e., a greater percentage of GM residue P was released under conventional tillage than under zero tillage in the first 2–10 weeks of residue placement. Wheat residues resulted in net P immobilization under zero tillage, but the amounts immobilized were less than 1 kg ha⁻¹. When net P mineralization occurred, the percentage of P released ranged from 24% of wheat P under conventional tillage to 74% of GM P under conventional tillage. The amounts of P released were 0.4 kg ha⁻¹ from wheat, 0.8 kg ha⁻¹ from canola, 0.4 kg ha⁻¹ from pea and 5.1–5.6 kg ha⁻¹ from clover GM residues. Therefore, only GM residues recycled agronomically significant amounts of P for use by subsequent crops in rotation. Phosphorus release was positively correlated with residue P concentration and negatively correlated with C/P and lignin/P ratios.     

Litterfall,decomposition, and nitrogen release in two age groups of trees in Casuarina equisetifolia plantations in the dry tropical Vindhyan plateau,India

Litterfall, decomposition, and N release in 5-year-old and 8-year-old plantations of Casuarina equisetifolia (Forst.) in the dry tropical region of the Vindhyan plateau were studied during 1989–1990. Maximum litterfall occurred in May. The total litterfall ranged from 7.2 to 9.9t ha^-1 year^-1 in the 5-year-old stand and from 11.3 to 12.7t ha^-1 year^-1 in the 8-year-old stand over the 2-year period. Photosynthetic branchlets contributed 87–95% to the total litter. The relative decomposition rates of litter components of the ash-free mass were highest in the rainy months (4.7 to 9.9mg g^-1 day^-1) followed by winter (2.8 to 3.6 mg g^-1 day^-1) and lowest in the summer months (1.7 to 3.0 mg g^-1 day^-1). Similar patterns were observed for N release. The annual decay constant was highest for cone litter and lowest for photosynthetic branchlets. During decomposition, the photosynthetic branchlets showed N immobilization in November and April, the twig litter in March, and the roots in January and February. N release per unit area (g m^-2) was maximum from the photosynthetic branchlets (5.3–6.3) followed by cones (4.4) > roots (3.4) > twigs (2.6–3.2). The combination of the litter C:N ratio, moisture, and temperature with the relative decomposition rate in a multiple regression analysis explained 66–84% of the variability in mass loss and 58–66% of the variability in N release.     

Yield and nitrogen response of lowland rice (Oryza sativa L.) to Sesbania rostrata and Aeschynomene afraspera green manure in different marginally productive soils in the Philippines

Organic-N fertilizers in the form of flood-tolerant, leguminous, stem-nodulating Sesbania rostrata and Aeschynomene afraspera may be useful alternatives to resource-poor rice farmers if applied as green manure. Therefore, the accumulation of N by these green manure species and their effect on the performance and yield of wetland rice (IR 64) was examined at four different sites in Luzon, Philippines. Soils deficient in N, P, and K were selected and compared with the fertile Maahas clay of the International Rice Research Institute (IRRI) at Los Baños. The green manure plants were grown under flooded conditions for 49 days in the wet season of 1987, chopped, and then ploughed in before transplanting rice seedlings. In a second experiment, the effect of S. rostrata green manure was studied under rainfed conditions. All green manure treatments were compared to an urea treatment (60 kg N ha^–1) and an untreated control. Both legumes developed well, even on the marginally productive soils. S. rostrata accumulated up to 190 kg N ha^–1 and A. afraspera even accumulated 196 kg N ha^–1 in the shoots. In all treatments, green manure increased grain yield significantly (P=0.05) over the untreated control, by 1.3–1.7 Mg ha^–1. The yields were comparable to those obtained with 60 kg N ha^–1 of urea fertilizer. S. rostrata caused the highest grain yield, of 6.5 Mg ha^–1 on the Maahas clay soil of IRRI. The apparent release of exchangeable NH ₄ ⁺ -N in the soils after green manuring and the rice grain yield response showed that both green manure species may provide sufficient available N throughout the development of IR 64 in the wet season. In the rainfed marginal soil site, green manure with S. rostrata produced even higher rice grain yields than urea. Green manure therefore seems particularly attractive for poor farmers on marginally productive soils, at least as a temporary strategy to improve yield and yield sustainability.     

Effects of nitrogen fertiliser and pesticide management on floodwater ecology in a wetland ricefield

We investigated the effects of N fertiliser and pesticide applications on the population dynamics of benthic molluscs in a tropical wetland rice field. Populations were monitored for two consecutive dry seasons in selected treatments during a study on the effects of agricultural practices on the floodwater ecology of tropical rice fields. The most abundant species recorded in the ricefields were the snailsMelanoides tuberculata andMelanoides granifera. Population densities and biomass values in planted plots ranged between 0 and 1530 individuals m^-2 and 0 and 1060 kg ha^-1, respectively. Snails were more abundant in unplanted than planted plots (1991: 170–2040 versus 0–1040 individuals m^-2, respectively). Populations in planted plots declined as the crop season progressed. Snail populations were significantly reduced by the broadcast application of mineral N fertiliser at 110 kg N ha^-1. There was little evidence that snails were affected by carbofuran or butachlor applications.     

Plant canopy effects on litter accumulation and soil microbial biomass in two temperate forests

The objective of this study was to determine whether differences in canopy structure and litter composition affect soil characteristics and microbial activity in oak versus mixed fir-beech stands. Mean litter biomass was greater in mixed fir-beech stands (51.9t ha⁻¹) compared to oak stands (15.7t ha⁻¹). Canopy leaf area was also significantly larger in mixed stands (1.96m² m⁻²) than in oak stands (1.73m² m⁻²). Soil organic carbon (C _org) and moisture were greater in mixed fir-beech stands, probably as a result of increased cover. Soil microbial biomass carbon (C _mic), nitrogen (N _mic), and total soil nitrogen (N _tot) increased slightly in the mixed stand, although this difference was not significant. Overall, mixed stands showed a higher mean C _org/N _tot ratio (22.73) compared to oak stands (16.39), indicating relatively low rate of carbon mineralization. In addition, the percentage of organic C present as C _mic in the surface soil decreased from 3.17% in the oak stand to 2.26% in the mixed stand, suggesting that fir-beech litter may be less suitable as a microbial substrate than oak litter.     

Above- and belowground ecosystem biomass and carbon pools in an age-sequence of temperate pine plantation forests

We assessed the successional development of above- and belowground ecosystem biomass and carbon (C) pools in an age-sequence of four White pine (Pinus strobus L.) plantation stands (2-, 15-, 30-, and 65-years-old) in Southern Ontario, Canada. Biomass and C stocks of above- and belowground live and dead tree biomass, understorey and forest ground vegetation, forest floor C (LFH-layer), and woody debris were determined from plot-level inventories and destructive tree sampling. Small root biomass (<5 mm) and mineral soil C stocks were estimated from soil cores. Aboveground tree biomass became the major ecosystem C pool with increasing age, reaching 0.5, 66, 92, and 176 t ha⁻¹ in the 2-, 15-, 30-, and 65-year-old stands, respectively. Tree root biomass increased from 0.1 to 10, 18, 38 t ha⁻¹ in the 2-, 15-, 30-, and 65-year-old stands, respectively, contributing considerably to the total ecosystem C in the three older stands. Forest floor C was 0.8, 7.5, 5.4, and 12.1 t C ha⁻¹ in the 2-, 15-, 30-, and 65-year-old stands, respectively, indicating an increase during the first two decades, but no further age-effect during the later growth phase. Mineral soil C was age-independent with 37.2, 33.9, 39.1, and 36.7 t C ha⁻¹ in the 2-, 15-, 30-, and 65-year-old stands, respectively. Aboveground ecosystem C increased with age from 3 to 40, 52, and 100 t C ha⁻¹ in the 2-, 15-, 30-, and 65-year-old stands, respectively, due to an increase in aboveground tree biomass. Belowground ecosystem C remained similiar in the early decades after establishment with 37, 39, and 39 t C ha⁻¹ in the 2-, 15-, and 30-year-old stands, but increased to 56 t C ha⁻¹ in the 65-year-old stand due to an increase in root biomass. The difference in total ecosystem C between the 2- and 65-year-old stand was 116 t C ha⁻¹. Our results highlight the importance of considering the successional development of forest ecosystem C pools, when estimating C sink potentials over their complete life cycle.     

Effects of earthworms on nitrogen mineralization

The influence of earthworms (Lumbricus terrestris and Aporrectodea tuberculata) on the rate of net N mineralization was studied, both in soil columns with intact soil structure (partly influenced by past earthworm activity) and in columns with sieved soil. Soil columns were collected from a well drained silt loam soil, and before the experiment all earthworms present were removed. Next, either new earthworms (at the rate of five earthworms per 1200 cm³, which was only slightly higher than field numbers and biomass) were added or they were left out. At five points in time, the columns were analyzed for NH ₄ ⁺ , NO ₃ ^– , and microbial biomass in separate samples from the upper and lower layers of the columns. N mineralization was estimated from these measurements. The total C and N content and the microbial biomass in the upper 5 cm of the intact soil columns was higher than in the lower layer. In the homogenized columns, the C and N content and the microbial biomass were equally divided over both layers. In all columns, the concentration of NH ₄ ⁺ was small at the start of the experiment and decreased over time. No earthworm effects on extractable NH ₄ ⁺ were observed. However, when earthworms were present, the concentration of NO ₃ ^– increased in both intact and homogenized cores. The microbial biomass content did not change significantly with time in any of the treatments. In both intact and homogenized soil, N mineralization increased when earthworms were present. Without earthworms, both type of cores mineralized comparable amounts of N, which indicates that mainly direct and indirect biological effects are responsible for the increase in mineralization in the presence of earthworms. The results of this study indicate that earthworm activity can result in considerable amounts of N being mineralized, up to 90 kg N ha^–1 year^–1, at the density used in this experiment.     

The earthworm population of a winter cereal field and its effects on soil and nitrogen turnover

The earthworm population in a winter cereal field in Ireland was studied over a 3-year-period and its effects on soil and N turnover were assessed. The mean annual population density was 346–471 individuals m^-2 and the mean biomass was 56.9–61.2 g m^-2. Twelve species were recorded, the most abundant being Allolobophora chlorotica followed by Aporrectodea caliginosa, and 242 mg at 5°C to 713 mg at 10°C in the case of juvenile Lumbricus terrestris. Gut contents (dry mass of soil) comprised 6.7–15.5% of the A. caliginosa live mass, and 9.7–14.7% of the Lumbricus terrestris mass. Annual soil egestion by the field population was estimated as 18–22 kg m^-2. Tissue production ranged from 81.7 to 218.5 g m^-2, while N turnover resulting from mortality was calculated as 1.5–3.9 g m^-2 depending on the year and the method of calculation. Earthworms were estimated to contribute an additional 3.4–4.1 g mineral N to the soil through excretion, mucus production, and soil ingestion. Independent estimates of N output via mucus and excretion derived from ¹⁵N laboratory studies with Lumbricus terrestris were 2.9–3.6 g m^-2 year^-1.