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Soil Biology & Biochemistry 35 (2003) 295302 www.elsevier.

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Effects of earthworm casts and compost on soil microbial activity and plant nutrient availability
Hala I. Chaoui1, Larry M. Zibilske2, Tsutomu Ohno*
Department of Plant, Soil and Environmental Sciences, 5722 Deering Hall, University of Maine, Orono, ME 04469-5722, USA Received 9 November 2001; received in revised form 6 September 2002; accepted 26 September 2002

Abstract Vermicomposting differs from conventional composting because the organic material is processed by the digestive systems of worms. The egested casts can be used to improve the fertility and physical characteristics of soil and potting media. In this study, the effects of earthworm casts (EW), conventional compost (CP) and NPK inorganic fertilizer (FT) amendments on N mineralization rates, microbial respiration, and microbial biomass were investigated in a laboratory incubation study. A bioassay with wheat (Triticum aestivium L.) was also conducted to assess the amendment effects on plant growth and nutrient uptake and to validate the nutrient release results from the incubation study. Both microbial respiration and biomass were signicantly greater in the CP treatment compared to EW treatment for the initial 35 days of incubation followed by similar respiration rates and biomass to the end of the study at 70 days of incubation. Soil NO2 increased rapidly in 3 the EW and CP treatments in the initial 30 days of incubation, attaining 290 and 400 mg N kg21 soil, respectively. Nitrate in the EW treatment then declined to 120 mg N kg21 soil by day 70, while nitrate in the CP treatment remained high. While ammonium levels decreased in the CP treatment as nitrate level increased with increasing incubation time, a low level of ammonium was maintained in the EW treatment throughout the incubation. The wheat bioassay study included two additional cast treatments (EW-N and EW2) to have treatments with higher levels of N input. Plants grown with CP or FT treatment had a lower shoot biomass and higher shoot N content than in EW-N and EW2 treatments, and also showed symptoms of salinity stress. Ionic strength and other salinity indicators in the earthworm cast treatments were much lower than in the CP treatment, indicating a lower risk of salinity stress in casts than in compost. All cast and compost amendments signicantly increased wheat P and K uptake compared to either the non-amended control or the mineral fertilizer treatment. The results show that casts are an efcient source of plant nutrients and that they are less likely to produce salinity stress in container as compared to compost and synthetic fertilizers. Published by Elsevier Science Ltd.
Keywords: Earthworm casts; N mineralization; Plant nutrient uptake; Microbial respiration; Vermicomposting

1. Introduction Vermicomposting is the digestion of organic materials by earthworms which produce excreta known as casts. Edwards (1995) reported that in a Rothamsted study with 25 types of vegetables, fruits or ornamentals, earthworm casts (EW) performed better than compost or commercial
* Corresponding author. Tel.: 1-207-581-2975; fax: 1-207-581-2999. E-mail addresses: ohno@maine.edu (T. Ohno), chaoui.1@osu.edu (H.I. Chaoui), lzibilske@weslaco.ars.usda.gov (L.M. Zibilske). 1 Department of Agricultural Engineering, The Ohio State University, 250 Agricultural Engineering Building, 590 Woody Hayes Drive, Columbus, OH 43210, USA. 2 USDA-ARS, Kika de la Garza Subtropical Agricultural Research Center, Integrated Farming and Natural Resources Research Unit, 2413 E. Hwy 83, Bldg 201, Weslaco, TX 78596-8344, USA. 0038-0717/03/$ - see front matter Published by Elsevier Science Ltd. PII: S 0 0 3 8 - 0 7 1 7 ( 0 2 ) 0 0 2 7 9 - 1

potting mixture amendments. It was suggested that the higher crop performance of the cast treatment was due to: better soil physical structure; presence of plant growth hormones; higher levels of soil enzymes; and greater microbial populations. The benecial effects of earthworm cast utilization in other horticulture settings have also been reported (Tomati et al., 1987; Hidalgo, 1999; Saciragic and Dzelilovic, 1986). EW typically have high N contents which suggests that they would be good sources of plant N (Parmelee and Crossley, 1988; Ruz-Jerez et al., 1992). Fresh casts often contain high ammonium levels, but rapid nitrication results in stable levels of both nitrogen forms due to organic matter protection in dry casts (Decaens et al., 1999). Nutrients in casts are initially physically protected, but this is reduced as the aggregate structure weakens over time (McInerney and

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Bolger, 2000). In addition to increased N availability, C, P, K, Ca and Mg availability in the casts is also greater than in the starting feed material (Orozco et al., 1996; Daniel and Anderson, 1992; Lavelle et al., 1992; Basker et al., 1993). Earthworm cast amendment has been shown to increase plant dry weight (Edwards, 1995; Lui et al., 1991) and plant N uptake (Zhao and Huang, 1988; Tomati et al., 1994). The benecial effect of EW has been observed in both horticultural plants (Tomati et al., 1987; Hidalgo, 1999; Saciragic and Dzelilovic, 1986) and in agronomic crops (Pashanasi et al., 1996). Cantanazaro et al. (1998) and Cox (1993) demonstrated the importance of the synchronization between nutrient release and plant uptake and showed that slower release fertilizers can increase plant yield and reduce nutrient leaching. EW could serve as a naturally produced slow release source of plant nutrients. Traditional composts also have agronomic value, but N immobilization (Sims, 1990), salinity effects (OBrien and Barker, 1996), and pathogen levels (Eastman, 1999) may be problematic. Vinceslas-Akpa and Loquet (1997) compared the effects of composting and vermicomposting lignocellulosic maple waste and reported that the vermicompost product had a lower C/N ratio, higher protein:organic C ratio, and higher levels of N, which indicates that the vermicompost products were more suitable for soil amendment use. In containerized production systems, EW used as an alternative soil amendment could help reduce several problems associated with the use of conventional synthetic fertilizer such as excessive leaching loss of nutrients and salinity-induced plant stress. In addition EW can improve soil porosity, and thus provide a better root growth medium. In this study, the effects of stabilized EW, compost and synthetic fertilizers on soil fertility and plant growth were investigated by determining mineralization rates of N, P, and K; microbial biomass-C levels; and microbial respiration in a laboratory incubation experiment. In addition, a greenhouse plant growth study with wheat (T. aestivium L.) was conducted to conrm the results of the incubation experiment.

Research Farm. The particular feedstock utilized for cast production and composting will inuence the specic chemical characteristics of the end products. However, we believe that the materials used in this study are representative of typical EW and compost available to growers. The compost and casts were stored at their native moisture state. The extractable NH-N and NO2-N in the soil and 4 3 amendment materials were determined by KCl extraction. Five g samples were extracted in 50 ml of 1 M KCl, placed on a reciprocal shaker for 15 min at 200 oscillations min21. The suspensions were ltered and analyzed for NH-N and 4 NO2-N using an autoanalyzer. Nutrient contents of the 3 amended soil mixtures were determined by extracting 5 g soil with 20 ml of modied-Morgan extract (1.25 M ammonium acetate, pH 4.8), shaking for 15 min at 18 oscillations min21, and ltration of the suspension (McIntosh, 1969). The P, K, Ca, and Mg content of the extract was determined by inductively coupled plasma atomic emission spectrometry (ICP-AES). The total carbon and nitrogen contents of the soil and amendment materials were determined using a LECO CN-2000 analyzer (St Joseph, MI). 2.2. Preparation of soilamendment mixtures This study was designed to evaluate the effect of earthworm cast and compost amendment on N release dynamics as compared to synthetic fertilizer. The treatments were: soil earthworm cast (EW); soil compost (CP); soil synthetic fertilizer (FT); and soil without amendment served as a control (CT). The treatment rates utilized were designed to equalize the quantity of N that would be available to the plant in a 28-day growth period. A preliminary incubation study estimated that the N mineralized in 28 days were 1.5% and 1.7 of the total nitrogen content in compost and casts, respectively. The amendment rates were calculated taking into account the N content, the bulk densities of the two amendments and the 530 cm3 volume of the pot. The EW treatment contained 267 g casts kg21 soil and the CP treatment contained 189 g compost kg21 soil. This resulted in an addition of 35% of compost and 36% of casts by volume, which minimized the effect of different bulk densities in the different treatments. These amendment rates would be appropriate for containerized production systems. The FT treatment received 63 mg N kg21 soil as (NH4)2SO4; 20 mg P kg21 soil as NaH2PO4H2O; and 108 mg K kg21 soil as K2SO4. All amendments were thoroughly mixed with the soil and placed in three separate units: an incubation jar to determine nutrient mineralization and microbial biomass, a respiration ask to determine microbial respiration, and a planted pot to bioassay N availability. The experimental design was a complete randomized design with three replications. Field capacity of the treatment mixtures as determined by gravimetric draining was 42% for the soil, 55% for the CP treatment (soil compost), 57% for the EW treatment

2. Materials and methods 2.1. Soils and materials characterization The surface horizon of a Nicholville (course-silty, mixed, frigid, Aquic Haplorthod) soil was obtained from the University of Maine Sustainable Agriculture Research Farm in Stillwater, Maine, USA. The soil contained 78% sand, 12% silt and 10% clay fractions as determined by the hydrometer method (Gee and Bauder, 1986). EW of Lumbricus rubellus were obtained from the Cape Cod Worm Farm (Buzzards Bay, Massachusetts, USA). Compost produced from cattle manure, leaves, and food scraps were obtained from the University of Maine Witter

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Table 1 Total C and N; 0.5 M K2SO4 extractable C; 1 M potassium chloride extractable NH-N and NO2-N; 1.25 M ammonium acetate (pH 4.8) extractable K and P; 4 3 and microbial biomass of the amendments and soil used in the study Material Total N Total C C/N ratio Extractable C Extractable NH-N 4 (mg kg21) (mg kg21) (g kg21) (g kg21) EW EW-N CP Soil 13.9 15.8 19.6 3.0 157 183 228 39 11.3 11.6 11.6 13.0 724 1510 5090 566 1.7 2.3 14.7 4.8 Extractable NO2-N 3 (mg kg21) 282 328 310 19 Extractable K Extractable P Microbial biomass (g kg21) (g kg21) (mg kg21) 1.93 2.29 11.4 0.27 3.71 4.31 2.42 0.01 658 148 3980 433

(soil EW), 77% for the EW-2 and 79% for the EW-N treatment. Soil moisture levels were maintained at their respective eld capacity levels by daily watering throughout the experiment. 2.3. Nitrogen mineralization study The open mineralization incubation pots were placed in a growth chamber kept at 70% relative humidity, 16 h of light at 20 8C and 8 h of dark at 16 8C. The mineralization jars were sampled on 3, 7, 15, 22, 28, 35, 43, 50, 57, and 70 days after incorporation by removing 60 g of soil. Nitrogen and other plant nutrient content was determined as described above. Microbial biomass was determined in the soils using a chloroform fumigation and extraction methodology (Voroney et al., 1991). Biomass C calculations followed Howarth and Paul (1994) using an extraction efciency constant of 0.35. 2.4. Microbial respiration study The microbial respiration asks were placed in a 19 8C incubator. The asks were sampled after 3, 7, 14, 22, 28, 35, 50, 57, 64, and 72 days of incubation. The alkali trap method was used to quantify the released CO2 (Landa and Fang, 1978). The stoppered 500 ml Erlenmeyer respiration asks contained soil treatments at eld capacity and scintillation vials containing 10 ml of 4 M NaOH. Flasks containing the alkali traps alone served as controls. The alkali traps were replaced at each sampling date and titrated with 1 M HCl (Stotzky, 1965). The respired CO2 was derived from titration data, corrected for the control. 2.5. Plant bioassay study Seeds of winter wheat (T. aestivium L.) were briey rinsed with 0.525% NaOCl solution, rinsed thoroughly with de-ionized water and germinated in a petri dish at 24 8C for 2 days prior to planting. Two seedlings were planted per pot and placed in a growth chamber set to identical conditions used in the incubation study above. Two treatments in addition to those used in the N mineralization study described above were added to the plant bioassay study: EW-2 which had an increased casts amendment rate (as

compared to EW) of 491 g casts kg21 soil (44% by volume), and EW-N at a rate of 330 g casts kg21 soil (36% by volume) which used a different cast lot which contained a higher total N content than EW was used (Table 1). On day 28, the plant shoots were harvested and washed. The plant matter was dried at 70 8C for 72 h, weighed and ground for elemental analysis. The N content was determined using the CN analyzer. The P and K contents were determined by dryashing the plant tissue at 450 8C for 5 h and re-dissolving the ash prior to analysis by ICP-AES. A saturated paste extract was prepared from soil sampled after plant harvest to determine soluble plant nutrients and electrical conductivity of the extract. The electrical conductivity values were converted to ionic strength using the regression reported by Grifn and Jurinak (1973).

3. Results 3.1. Amendment materials Although the compost material has a higher absolute N and C content than the casts, the C/N ratios were very similar (Table 1). The compost material had a higher level of extractable NH than the casts, but both contained 4 comparable amounts of NO2 (Table 1). The lower levels of 3 NH found in the EW are probably due to the high 4 nitrication rates associated with cast stabilization (Decaens et al., 1999). The compost contained a greater amount of extractable K and lower amount of extractable P than the EW casts. 3.2. Microbial respiration and biomass The average daily CO2 production is shown in Table 2. The elevated respiration across all treatments at day 3 is most likely due to the stimulation of the soil microbial activity by the greater oxygen availability attributable to physical mixing of the soil and amendments at the start of the experiment. Respiration in the control was relatively stable from day 7 to the end of the experiment. As shown in Table 2, respiration levels for the CP treatment were signicantly higher than in the EW treatment for the initial 35 days and they then became statistically equivalent until

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Table 2 Average soil respiration in microgram of CO2 produced per gram of dry soil per day in the earthworm cast (EW), compost (CP), fertilizer (FT) and unamended control (CT) treatment soils during the incubation Days 3 7 14 22 28 35 50 57 64 72
a

EW (mg CO2 g soil21 day21) 187 ba 63 b 86 b 44 b 77 b 82 b 117 a 71 a 79 a 66 a

CP (mg CO2 g soil21 day21) 464 a 324 a 227 a 186 a 190 a 155 a 122 a 76 a 89 a 77 a

FT (mg CO2 g soil21 day21) 115 b 39 b 47 c 39 b 23 c 21 c 14 b NDb 24 b 4b

CT (mg CO2 g soil21 day21) 138 b 35 b 52 c 48 b 32 c 25 c 16 b 14 b 12 b 6b

Fishers protected mean separation test was used at the p , 0.05 level within each treatment. Means within a row followed by the same letter are not signicantly different. b ND, not determined.

the end of the incubation. In general, microbial biomass increased rapidly during incubation, peaking at 15 days for both casts-treated (EW) and compost treated (CP) soils (Table 3). The level of microbial biomass was signicantly lower in the EW treatment than in the CP treatment in the initial 28 days which was probably due to the initially lower microbial biomass contributed by the casts (Table 1). 3.3. Nutrient mineralization The quantity of KCl-extractable NH-N decreased 4 steadily during the entire incubation time in the FT treatment which received (NH4)2SO4 and decreased rapidly within the initial 15 days in the CP treatment (Fig. 1A). The KCl-extractable NO2-N levels generally increased with 3 incubation time which was probably due to the transformation of ammonium to nitrate (Fig. 1B). The Morgan soil test levels of both P and K did not change during the incubation which suggests that the content of these elements in the amendments was sufcient for microbial growth. Extractable P (mg P kg21 soil) in the treatments were: EW, 535; CP, 245; FT, 6.1; and CT, 5.1. Extractable K (mg K kg21 soil) in the treatments were: EW, 810; CP, 2830; FT, 1020; and CT, 380. 3.4. Plant bioassay analysis Plant dry shoot biomass data are shown in Fig. 2. The EW-N, EW-2 and CP shoot weights were signicantly greater than in the EW treatment which is probably due to higher levels of plant available N in these treatments. The EW treatment biomass was statistically equivalent to the NPK FT treatment and all amendment treatments signicantly increased biomass over the unamended control treatment. The shoot N content (dry shoot weight X N concentration) was the highest for the NPK FT treatment, followed by the CP treatment (Fig. 3). The EW-2 and EW-N

treatments had shoot N contents which were statistically equivalent followed by the EW treatment which has signicantly lower N uptake than EW-2, but not EW-N (Fig. 3). Shoot P content in all the EW and CP treatments were higher than in the FT and CT treatments, demonstrating that these amendments may be adequate sources of P (Fig. 4A). The K content results were similar with uptake from the EW-2, EW-N, and CP treatments being higher than from the other treatments (Fig. 4B).

4. Discussion The elemental composition of the EW and compost materials used suggests that the materials have potential as alternative plant nutrient sources. The low C/N ratio indicates that the casts and compost would be effective sources of N through rapid N mineralization reactions
Table 3 Microbial biomass in microgram of C per gram of dry soil per day in the earthworm cast (EW), compost (CP), fertilizer (FT), and unamended control (CT) treatment soils during the incubation CP FT CT Days EW (mg C g soil21) (mg C g soil21) (mg C g soil21) (mg C g soil21) 3 7 15 22 28 35 43 50 57 70
a

350 ba 634 b 653 b 284 bc 345 b 297 ab 404 a 326 b 424 a 397 b

730 a 677 a 973 a 639 a 688 a 495 a 316 a 589 a 318 ab 866 a

140 c 157 b 404 c 433 abc 358 ab 316 ab 120 b 187 b 178 b 121 c

183 c 207 b 175 d 179 c 191 b 166 b 283 a 357 b 254 b 272 bc

Fishers protected mean separation test was used at the p , 0.05 level within each treatment. Means within a row followed by the same letter are not signicantly different.

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Fig. 3. Dry matter biomass of shoots in the earthworm cast (EW), the EW cast at higher amendment rate (EW-2), earthworm cast with a higher native N content (EW-N), compost (CP), synthetic NPK fertilizer (FT), and control (CT) treatment pots. Fishers protected mean separation test was used at the p , 0.05 level between each treatment mean. Means within a row followed by the same letter are not signicantly different.

Fig. 1. Potassium chloride (1 mol l21) extractable levels of (A) ammonium nitrogen and (B) nitrate nitrogen during the incubation period in EW, compost (CP), synthetic NPK fertilizer (FT), and control (CT) treatments.

(Tisdale et al., 1993). The compost material contained a much higher level of soluble organic C (K2SO4-extractable) than the cast materials. This is indicative of the lesser degree of decomposition that the compost has undergone and suggests that the material is still rich in labile carbon compounds which can serve as an energy source for

Fig. 2. Total potassium chloride (1 mol l21) extractable (ammonium plus nitrate nitrogen) levels during the incubation period in EW, compost (CP), synthetic NPK fertilizer (FT), and control (CT) treatments.

Fig. 4. Plant shoot N uptake in the earthworm cast (EW), the EW cast at higher amendment rate (EW-2), earthworm cast with a higher native N content (EW-N), compost (CP), synthetic NPK fertilizer (FT), and control (CT) treatment pots. Fishers protected mean separation test was used at the p , 0.05 level between each treatment mean. Means within a row followed by the same letter are not signicantly different.

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microbes. Casts are a byproduct of the digestion process so they would be expected to be lower in soluble organic compounds which are used as a microbial energy substrate. Although the nutrient content in casts and composts are much lower than that found in synthetic fertilizers, they are comparable in nutrient content to that typically found in other secondary sources such as animal manure (Troeh and Thompson, 1993). As with other carbon-rich amendment materials, casts and compost have the potential to increase soil organic matter levels and improve soil quality. The quantity of soluble C was 6.3 times greater in the CP material than in the EW method and suggests that the initial microbial activity is linked to the level of soluble C in the treatments (Table 1). Likewise, the microbial activity in the later stages of the incubation may have been controlled by the nearly equivalent amount of total of C added to the soil (CP, 25.1 g C added; EW, 21.0 g C added). There were no signicant differences in respiration rates between the soil amended with mineral fertilizer (FT) and the control (CT), suggesting that microbial activity was not limited by inadequate nutrient levels in the soil. In addition to the C status in controlling microbial respiration, improved aeration of the EW and CP treatment are also thought to be involved. The greater pore volume in cast and compost amended soils increases the availability of both water and nutrients to microorganisms in soils (Scott et al., 1996). EW may have reduced levels of microbial biomass due the earthworm use of microbes as an energy source (Bohlen and Edwards, 1995). Microbial biomass after 35 days of incubation was highly variable for the EW and CP treatments, increasing and decreasing with incubation time. Microbial biomass in the EW treatments did not signicantly differ from the control (CT) in the four nal (Days 43, 50, 57 and 70) sampling dates (Table 3). However, soil respiration rates for the EW treatment was signicantly higher than in the control soil in these nal four sampling dates (Table 2). The higher respiration rates in the EW treatment in conjunction with no difference in biomass as compared to the control could be due to the presence of different classes of microorganisms in the casts which might have a different respiration to biomass ratio than the organisms found in the soil, such as the fungi observed by Marinissen and Dexter (1990). Bohlen and Edwards (1995)

Fig. 5. Plant shoot (A) P content and (B) K content in the earthworm cast (EW), the EW cast at higher amendment rate (EW-2), earthworm cast with a higher native N content (EW-N), compost (CP), synthetic NPK fertilizer (FT), and control (CT) treatment pots. Signicant differences at the 5% level using Duncans multiple range test between treatment means are indicated by differing letters. Means within a row followed by the same letter are not signicantly different.

and Daniel and Anderson (1992) reported similar differences between soil microbial biomass and respiration level. This incubation study suggests that EW could serve as a naturally produced, slow release source of plant nutrients. The slope of NO3-N production over time shown in Fig. 1B corresponds to an average rate of the microbially mediated nitrication reaction (mg NO2-N kg21 soil day21). The 3 greater nitrication in the initial 28 days for the CP treatment (12.1 ^ 0.7 mg NO2-N kg21 soil day21) than in 3 the EW treatment (4.7 ^ 0.3 mg NO2-N kg21 soil day21) 3 which was signicantly different using the t-test to evaluate regression slopes (Zar, 1984) suggests that this microbially mediated process was higher in CP than in EW treatment. This may be due to the organic matter being more readily available to microorganisms in compost than in the casts since organic matter in casts is thought to be stabilized by the formation of a clay casing (Chan and Heenan, 1995; Shipitalo and Protz, 1989). The release of total extractable N (ammonium nitrate) in the EW and CP treatments compared with the NPK FT treatment is shown in Fig. 5. The total extractable N

Table 4 Chemical characterization of the saturated paste extracts of the soils from the plant bioassay study at harvest Days Nitrate-N Phosphorus Potassium Sodium Ionic Strength EW (mmol l21) 0.87 ca 0.14 c 2.0 c 2.3 c 18.2 c EW-2 (mmol l21) 0.01 d 0.20 a 1.8 c 2.2 c 15.0 cd EW-N (mmol l21) 0.04 d 0.17 b 1.9 c 2.1 c 14.6 d CP (mmol l21) 2.37 b 0.13 c 21.3 a 17.2 a 66.1 a FT (mmol l21) 3.44 a 0.03 d 12.1 b 7.9 b 62.5 b

a Fishers protected mean separation test was used at the p , 0.05 level within each treatment. Means within a row followed by the same letter are not signicantly different.

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peaked at 35 days for the EW treatment and at 43 days for the CP treatment while the FT levels were highest at the start of the incubation and declined with incubation time. The decrease in nitrate and in extractable total N after the rst 7 weeks is probably due to denitrication. The levels of P and K found in the EW and CP treatments reected the quantity of the nutrients present in the amendment materials (Table 1). These results show that both EW and CP are good supplemental sources of readily available P and K, as well as for N. The plant study was conducted to test whether the results and interpretations of the incubation studies were supported using a bioassay. There was evidence of salinity stress in the CP and FT plants with some shoots displaying leaf tip burn. The saturated paste soil extract analysis conducted at harvest indicated that the levels of NO2-N and ionic strength in the CP and FT were much 3 higher than in the earthworm cast treatments suggests that the burned leaf tips in the CP and FT plants at harvest were symptoms of salinity stress (Table 4). Nitrate-N in the EW-2 and EW-N extracts was less than 2% of that found in CP (Table 4), although the initial extractable N levels in EW-N and EW-2 were similar to that of compost (Table 1). This indicates a slower N release in casts and a possible lower risk of nitrate leaching with the use of casts as compared to compost. The CP treatment contained the highest quantity of K and Na in the saturated paste extract which may have also contributed to the salinity stress in that treatment (Table 4). This result suggests that casts may be safer than compost and water-soluble synthetic fertilizers in containerized systems. In summary, when sufcient quantities of casts (EW-2) and casts higher in N content (EW-N) were used to provide sufcient N to the plant, dry shoot biomass was greater than the yield obtained with equivalent quantity of NPK fertilizer and statistically equivalent to a treatment where the N source was compost. Shoot N content was higher in the CP and FT treatments than in any of the earthworm cast treatments and evidence of salinity stress symptoms were observed. These results suggest that EW in EW-2 and EWN resulted in higher plant biomass production due to a slower rate of nitrogen mineralization that was more synchronized with the plant requirements (Cox, 1993). Ionic strength and other salinity indicators in earthworm cast treatment were much lower than in the CP treatment, indicating that the casts used in this study did not cause adverse salinity stress. The plant biomass and shoot elemental content data show that casts are an efcient source of plant nutrients and that the slower rate of N release in EW gives it an advantage as compared to compost and synthetic fertilizers. The nutrient content of the organic waste fed to earthworms determines the level of nutrients present in the obtained casts (Lavelle et al., 1992), and compost is also affected similarly by the raw material used. However, comparable results are expected regardless of the

specic source of casts with respect to the physical structure resulting from the gut digestive processes which is responsible for the general slow nutrient release characteristic of EW (Shipitalo and Protz, 1986).

Acknowledgements Support for this work was provided by Hatch funds from the Maine Agricultural and Forest Experiment Station. MAFES Journal Publication 2603.

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