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Published in Soil Sci. Soc. Am. J. 68:538-544 (2004).
© 2004 Soil Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA

DIVISION S-4—SOIL FERTILITY & PLANT NUTRITION

Nitrogen Use Efficiency of 15N-labeled Poultry Manure

Ingrid K. Thomsen*

Danish Inst. of Agric. Sci., Dep. of Agroecology, Research Centre Foulum, P.O. Box 50, DK-8830 Tjele, Denmark

* Corresponding author (ingrid.thomsen{at}agrsci.dk).


    ABSTRACT
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
The timing between poultry manure N mineralization and crop N uptake as influenced by manure application time (winter or spring) and bedding material [barley (Hordeum vulgare L.) straw, wood chips, or no bedding] was investigated with 15N-labeled poultry manure. Reference plots received either a high or low dose of 15N-labeled mineral fertilizer in spring or were without manure and fertilizer N. The 15N recovery was measured for 1 yr in spring barley and for the two following years in perennial ryegrass (Lolium perenne L.). Barley grain yield and N uptake were significantly higher when poultry manure had been applied in spring compared with application in winter. The barley 15N recovery was 15% for winter-applied manure and 38% for manure applied in spring. The recovery of manure 15N from spring-applied poultry manure corresponded to 82% of what was taken up by barley in the fertilizer-amended soil, where 15N recovery averaged 46%. Bedding in poultry manure had no effect on crop 15N uptake. Ryegrass recoveries of 15N from manure and fertilizer were 4 to 6% in the second year and 1 to 2% in the third year. After 3 yr, 94% of 15N of spring-applied poultry manure and 88% of mineral fertilizer were recovered in crops plus soil. Total recovery of 15N from winter-applied poultry manure averaged 56%. The bedding had no influence on total 15N recovery. It is concluded that although poultry manure contains bedding with high C:N ratios which potentially could immobilize N for a period, optimum time for manure application is in the spring.

Abbreviations: CAN, calcium ammonium nitrate • MFE, mineral fertilizer equivalent


    INTRODUCTION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
NUTRIENT CONTENTS OF poultry manures are among the highest of all animal manures, and the use of poultry manure as soil amendment for agricultural crops will provide appreciable quantities of all important plant nutrients (Sims and Wolf, 1994). Ammonium-N (NH4–N) is a significant part of total N in poultry manure, which additionally contains uric acid. Uric acid metabolizes rapidly to NH4–N in most soils, and the net result of the high NH4–N and uric acid contents in poultry waste is that a large percentage of N can be converted to nitrate-N (NO3–N) within a few weeks (Sims and Wolf, 1994). This may increase the risk of NO3–N leaching from poultry manure amended soils unless manure is applied at a time that matches crop N uptake.

To absorb moisture in poultry houses, bedding materials such as straw, sawdust, or wood chips may be used. The carbonaceous bedding mixed into the manure is subsequently a part of the manure and may affect nutrient transformations both during storage and after application to the field. The influence of bedding during storage may relate to both a modifying effect on the physical conditions as regards to air and water transport in a pile and by causing N immobilization due to the high C:N ratios of the bedding (Mahimairaja et al., 1994). Bedding material may also influence crop N availability. Kirchmann (1989) mixed increasing amounts of cereal straw into poultry manure and found that crop N uptake depended on the C:N ratio obtained. Martín-Olmedo and Rees (1999) incubated cellulose together with poultry manure and they likewise found that different amounts of cellulose resulted in increased immobilization.

Nitrogen immobilization in the field caused by manure containing bedding has various impacts. Immobilization may diminish the risk of NO3–N leaching if manure is applied in autumn or winter where leaching losses would otherwise occur due to either low or missing crop N uptake. On the other hand, if manure is applied immediately before the growing season, N immobilization may reduce the amount of plant available N in soil for a period.

The crop utilization of poultry manure can be investigated by either the difference method, that is, soil with and without manure, or by the use of 15N-labeled manure. The production of 15N poultry manure is feasible, and quite homogeneous manure can be obtained (Uenosono et al., 2002). An advantage of 15N-labeled manure use is the higher accuracy of the determination of the residual effect of poultry manure, which may be difficult to detect except where high application rates have been used (Chambers et al., 1996).

The objective of this study was to investigate the optimization of the timing between N mineralization from poultry manure and crop N uptake patterns. Different application times were tested for 15N-labeled poultry manures without bedding and manures containing bedding which potentially could change the N immobilization and mineralization processes in soil.


    MATERIALS AND METHODS
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
Production of Poultry Manure
Nitrogen-15 labeled grains of spring barley containing 1.82% N in dry matter (7.779 atom% 15N) and field peas (Pisum sativum L.) with 4.00% N in dry matter (4.939 atom% 15N) were milled (4 mm). Six chickens (Gallus domesticus) were fed for 20 d with daily portions of 35 g peas mixed with 85 g barley. The chickens were fed with unlabeled barley and peas for the first 5 d and with the 15N-labeled feed for the remaining days. The manure produced was collected two times per day and frozen after feathers and other impurities had been removed. The average manure production per chicken was 139 g d–1 with a dry matter content of 22%. Subsamples of the manure from Days 1, 4, 7, 10, 13, 16, and 19 were collected for analysis.

On Day 13, when the chickens had been fed with 15N-labeled feed for 7 d, the content of 15N in the manure had reached a sufficiently stabilized level (Fig. 1) . The manure portions from Day 13 and onwards were accordingly mixed and divided into three portions. One of the manure portions was mixed with unlabeled barley straw (1.4% N, 44.6% C); one with wood chips (0.2% N, 48.5% C), and one remained without bedding. The ratio between bedding and manure was based on a daily use of 5 g of bedding per chicken, giving a manure-to-bedding dry-matter ratio of 1.0:0.2. The 15N-content of the feed was 6.430 atom% 15N while the 15N-content of total N in the manures averaged 4.180 atom% 15N (Table 1). The three manures were stored at 13 to 15°C. After 10 d, the manures were weighed into portions and frozen. Sufficient portions were prepared to allow for two application times in two successive field experiments. Characteristics of the manure are shown in Table 1.



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Fig. 1. Atom% 15N of NH4–N and total N in poultry manure during the feeding period. The feed was 15N-labeled from Day 6 and onward. Standard errors (n = 6) are indicated.

 

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Table 1. Characteristics of manures after 10 d of storage.

 
Soil and Site
Polyvinyl chloride cylinders (30-cm diam., 30-cm length) were inserted into the soil in a field at the Research Centre Foulum in Denmark (56°30' N, 09°35' E). One insertion took place in autumn 1998, where 27 cylinders were placed in three rows (blocks) of nine with a distance of 2 m. Another insertion took place in autumn 1999, approximately 50 m apart. The cylinders protruded 5 cm above the soil surface, and the soil within the cylinders was level with the surrounding soil surface. The soil is a loamy sand (Typic Hapludult, mixed, mesic) with 72 g clay and 111 g silt kg–1 dry soil. Soil C and N contents are 17.3 g and 1.5 g kg–1 dry soil, respectively. Annual mean temperature at the site is 7.3°C, the highest and lowest being 15.4°C (July) and –0.5°C (January, February). Annual mean precipitation is 626 mm.

Field Experiment
Application of the poultry manure took place in winter and in spring. The soil inside each cylinder was removed to a 15-cm depth and sieved (1 cm). A portion of the manure (1.2 kg m–2 wet weight) was mixed with the sieved soil and the soil-manure mixture returned to the cylinder. The manure supplied 19.0 (manure with straw), 18.5 (manure with wood chips), and 19.1 (manure without bedding) g total N m–2, respectively. Unmanured cylinders received either a high (10 g N m–2) or low (5 g N m–2) dose of 15NH415NO3 (5.0 atom% 15N) in spring or were held without N. All cylinders received P (4.5 g m–2) and K (5.5 g m–2). See Table 2 for details on dates for the field operations. The cylinders and the area surrounding the cylinders were sown to spring barley and perennial ryegrass in the first growth season. The barley was harvested at maturity, and the ryegrass was cut one time the following autumn. In spring before the first ryegrass cut in the second and third year, the ryegrass was fertilized with calcium ammonium nitrate (CAN; 3.8 g N m–2), P (2.8 g m–2), and K (13.9 g m–2). The ryegrass was cut three times in the second and third year. After the first and second cut of the ryegrass, cylinders received 2.0 g N m–2 as CAN. Cylinders that received no manure or fertilizer N at the beginning of the field trial (no N) remained without N and were supplied only with P and K in the 3 yr. After the last cut in the third experimental year, soil was sampled from each cylinder by pooling six soil cores (20-cm soil depth) from each cylinder.


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Table 2. Details on the field experiments.

 
All treatments were performed in three replicates. Two identical experiments with an interval of 1 yr were performed and are referred to as the first and second series, respectively (Table 2).

Analyses
Total N in the manures was determined after Kjeldahl digestion with a Kjeltec Auto 1030 system (Tecator, Höganäs, Sweden). Nitrate-N and NH4–N were extracted in 2 M KCl for 1 h with a ratio between manure (wet weight) and KCl of 1:50. The NO3–N and NH4–N in the KCl extracts were analyzed on a Technicon Autoanalyzer II (Bran+Luebbe GmbH, Norderstedt, Germany). The 15N in KCl extracts and Kjeldahl digests was determined by ANCA-MS (Europa Scientific, UK) after diffusion (Brooks et al., 1989). Manure dry matter content was determined on subsamples after drying at 80°C for 24 h. Manure ash content was determined by weighing before and after ignition at 525°C for 6 h. Total manure C was determined after dry combustion on a Leco model 521-275 (Leco Corporation, Svenska AB, Upplands Väsby, Sweden). Losses of total N and C were estimated relative to the ash content on the assumption that ash was conserved in the manure during the 10 d of storage.

The spring barley was separated into grain and straw with a plot threshing machine. The plant materials were dried at 80°C for 24 h and ground in a centrifugal mill (Retsch Ultra-Centrifugal Mill, Type ZM 1, Haan, Germany). Subsamples were further ball milled for 20 min (Retsch Ball Mill, Type S 1). Soil was dried (80°C) for at least 48 h and ball milled for 20 min. The finely milled plant and soil samples were packed into tin capsules and analyzed for total N and 15N by ANCA-MS (Europa Scientific, UK). The 15N contents were corrected for natural background 15N abundance.

The availability of 15N in poultry manure was related to the availability of mineral fertilizer 15N by calculating the mineral fertilizer equivalent (MFE): MFE = (% uptake of 15N from manure) x 100/(% uptake of 15N from mineral fertilizer) (Christensen, 1996). Data were analyzed statistically by the GLM procedure in the SAS statistical analysis system (SAS Institute, 1988). Significant effects were separated by Duncan's test at P = 0.05.


    RESULTS AND DISCUSSION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
Manure Preparation
During the 10 d of storage, about 20% of the initial C content was lost from the three manures, whereas 7 to 10% of initial N content was lost. The mixing with bedding did not influence C and N losses. The C losses could mainly be ascribed to chemical processes in the conversion of uric acid to ammonium rather than to biological processes. The weight of each of the three manure portions was approximately 2 kg and the small portions prevented composting from taking place in the relatively short storage period.

In a 12-wk incubation study, Mahimairaja et al. (1994) found that bedding materials reduced N losses, with wheat straw and peat being superior to other tested materials. The straw amendment reduced N losses from 17 to 11% (Mahimairaja et al., 1994). Nitrogen losses during composting may, however, account for much higher proportions. More than half of the total N content in poultry manure was lost during composting in the studies of Tiquia and Tam (2000) and Uenosono et al. (2002). As the N lost by storage would mainly be from the mineral N pool, high losses during storage will influence the fertilizer value of the manure as a relatively larger proportion of N subsequently will be present in the organic pool.

Yield and Nitrogen Uptake after Application of Poultry Manure and Mineral Fertilizer
The amount of N applied as poultry manure to the first barley crop averaged 19 g N m–2 while the high and low doses of mineral fertilizer were 10 and 5 g N m–2, respectively. It was estimated that yield of spring barley supplied with poultry manure in spring would be in the range between yields obtained with the two doses in mineral fertilizer. However, yields were higher after the spring-applied manure than with the highest mineral-fertilizer application (Fig. 2) . In the first series, spring manured barley grain yield averaged 839 g m–2, whereas the barley grown with the high and low dose of mineral fertilizer yielded 695 and 500 g m–2 grain dry matter, respectively. Yield was significantly lower (P < 0.05) when poultry manure was applied in winter, where barley in the first series yielded 475 g grain dry matter m–2 (Fig. 2). The yield obtained after winter application of the poultry manure was similar to yield of barley supplied with the low dose of mineral fertilizer in spring. The barley response to poultry manure was the same whether or not the manure had been mixed with bedding. Barley grain yield was generally lower in the second series, but the relationships between the different treatments were the same as in the first series (Fig. 2).



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Fig. 2. Grain yield of spring barley in the first growth season after application of poultry manure (black bars), mineral fertilizer (gray bars), or no N (white bars). Standard errors (n = 3) are indicated. Bars denoted with the same letter are not significantly different (P = 0.05).

 
The ryegrass underseeded in barley was harvested in the year of sowing and three times in each of the two succeeding years. All ryegrass plots, except for the plots previously held without N at the beginning of the field trial, received mineral fertilizer in the last 2 yr. In general, the ryegrass held without N yielded less than the ryegrass receiving N (Fig. 3) . There were, however, no differences between yield of ryegrass grown after manure application and yield of ryegrass only supplied with mineral fertilizer. The lack of residual value of the manure in ryegrass yield was observed in both series, and relations were the same whether yields from single years were compared or the total cumulative yield was considered.



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Fig. 3. Dry matter yield of ryegrass grown after application of poultry manure, mineral fertilizer, and no N. Bars are divided into ryegrass harvested in the year of sowing (black) and cumulative ryegrass yield of the three cuts in the second (gray) and third (white) year after initiation of the experiment. Standard errors are shown for the cumulative yield (n = 3). Bars denoted with the same letter are not significantly different (P = 0.05).

 
The N uptake in barley reflected the variations found for grain yield. The highest N uptake was found in the first experimental series where N corresponding to 16 g N m–2 was taken up by the barley supplied with poultry manure in spring (Fig. 4) . When the poultry manure was applied in winter, about 10 g N m–2 was taken up. In the second experimental series, the uptake after the previous winter and spring application was 7 and 11 g N m–2, respectively (Fig. 4). Thus, for both barley crops, the total N uptake after winter application of poultry manure corresponded to only 65% of the N uptake after application of the manure in spring. Similar amounts of N were taken up by barley supplied with manure mixed with bedding and barley receiving manure containing no bedding. The N uptake in the barley supplied with poultry manure in winter was in both years of the same size as in barley receiving the low dose (5 g N m–2) of N in mineral fertilizer in spring. When applied in spring, N uptake in the manured plots, which had received about 19 g N m–2, exceeded what was taken up in the barley supplied with the high dose (10 g N m–2) of mineral fertilizer (Fig. 4).



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Fig. 4. Nitrogen uptake in grain (black bar) and straw (white bar) of spring barley in the first growth season after application of poultry manure, mineral fertilizer, and no N. Standard errors are shown for the cumulative N uptake in grain and straw (n = 3). Bars denoted with the same letter are not significantly different (P = 0.05).

 
As with dry matter yield, N uptake in the ryegrass grown in the 2 yr following barley harvest was rather similar for plots previously supplied with manure and plots only supplied with mineral fertilizer (Fig. 5) . No residual effect of the poultry manure was detected in ryegrass total N uptake when compared with ryegrass that had received mineral fertilizer in the high or low dose. Likewise, Chambers et al. (1996) found a low residual N value of poultry manure when applied at agronomically sensible rates. Only the ryegrass held without N in the 3 yr differed from the other plots by having a significantly lower cumulative N uptake (Fig. 5).



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Fig. 5. Nitrogen uptake in ryegrass grown after application of poultry manure, mineral fertilizer, and no N. Bars are divided into N uptake in ryegrass harvested in the year of sowing (black) and cumulative N uptake in ryegrass in the three cuts in the second (gray) and third (white) year after initiation of the experiment. Standard errors are shown for the cumulative N uptake (n = 3). Bars denoted with the same letter are not significantly different (P = 0.05).

 
Crop Nitrogen-15 Recovery
The recovery of 15N illustrates very clearly the importance of application time for poultry manure (Fig. 6 , Table 3). After the winter application, about 15% of the 15N in the poultry manure was taken up by the first barley crop. When the application of the manure took place in spring, the uptake of 15N was about two or three times higher, corresponding to 43 and 33% of manure 15N in the first and second series, respectively. The mixing of bedding with manure did not affect the barley 15N uptake neither at the winter nor the spring application.



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Fig. 6. Cumulative 15N recovery in spring barley (grain and straw) and ryegrass 3 yr after application of 15N-labeled poultry manure and mineral fertilizer. Standard errors (n = 3) are indicated.

 

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Table 3. Cumulative 15N recovery in crops and soil after three growing seasons.

 
The barley uptake of 15N from the mineral fertilizer was almost identical for the high and low dose (Fig. 6). The barley crop grown in 1999 took up 52% of applied fertilizer 15N when averaged across the two doses, whereas a lower amount, corresponding to 40% of the applied 15N, was taken up in 2000. The MFE (Christensen, 1996) gives an indication of the utilization of organic manure N compared with mineral fertilizer N and enables comparisons of the 15N availability for different manures or other organic residues in different years and on different soils. It appears that MFE for spring-applied poultry manure was identical in the 2 yr although the actual 15N uptake differed. The averaged MFE of the spring-applied poultry manures was 82 in both years. The MFE for the three poultry manures applied in winter averaged 29 in the first barley crop (1999) and 35 in the barley crop grown in the second series (2000).

Sørensen and Jensen (1998) estimated MFE for several experiments with 15N-labeled ruminant feces and urine. They found that MFE for feces ranged from 13 to 33. The MFE of 82 for the spring-applied poultry manure was thus very high when compared with other types of solid manure, and was of the same size as obtained for ruminant urine (Sørensen and Jensen 1998). Kirchmann (1989) found that wheat recovered 28% of 15N in poultry manure without bedding and from 5 to 31% when bedding (oat straw) had been mixed with the manure. From the results of Kirchmann (1989), Christensen (1996) calculated MFE in the range of 9 to 56 for the poultry manures depending on bedding content. The three manures had C:N ratios of 6 to 7:1 (Table 1). Christensen (1996) estimated a MFE of 51 for a poultry manure with a comparable C:N ratio of 8 used in Kirchmann (1989). The higher MFE obtained for spring-applied manure in the present experiment may have been caused by lower gaseous N losses during preparation of the manures. The poultry manure in the Kirchmann (1989) study was air-dried, a process which may have promoted considerable N losses. Sistani et al. (2001) compared four methods of poultry manure drying, including freeze-drying, with no drying before analysis. They found that 21 to 27% of total N was lost irrespective of drying method. Also, Giddens and Rao (1975) found high N losses on drying but found that air-drying resulted in greater N loss than rapid drying with heat. As a more recalcitrant part of N will be left in manures that have been dried, lower plant N availability can be expected than in the present experiment where no drying was done.

In the ryegrass harvested in the first autumn after the spring barley harvest, a further 1 to 2% was taken up of the poultry manure 15N (Fig. 6). The uptake of 15N applied in mineral fertilizer was of the same size.

In the two growth seasons following the year of application, 15N uptake was determined in cuts of ryegrass. The almost-parallel lines in Fig. 6 indicate that the 15N uptake was almost independent of the origin of the applied 15N. In the second year, similar amounts (3–4%) of 15N were taken up by ryegrass on plots amended with poultry manure applied in winter and plots supplied with the two doses of mineral fertilizer in spring. Slightly more 15N (4–6%) was recovered in ryegrass grown after spring application of poultry manure. In the third year, an additional 1 to 2% 15N of the initial manure and fertilizer was taken up. The higher 15N recovery in the second year ryegrass grown after previous spring application of poultry manure resulted in a cumulative 15N recovery of 9% in both series (Table 3). Ryegrass recovery of 15N from manure applied in winter and from mineral fertilizer was 6%.

As different amounts of N were applied with the poultry manures and the two doses of mineral fertilizer, the almost-similar percentages for second and third year 15N recovery imply that the actual amounts of N taken up by the crops differed in the 2 yr. However, the variations in 15N uptake were too small to result in significant differences in total N uptake between ryegrass grown after an application of poultry manure and ryegrass grown after the application of mineral fertilizer (Fig. 3).

The shapes of the curves in Fig. 6 are very similar to 15N recoveries for solid ruminant manure shown by Jensen et al. (1999) and for the pig slurry examined in Sørensen and Amato (2002). The curves indicate that the type of N source and the application time determine first-year crop N uptake, whereas the relative N uptake in the following years is almost independent of the origin of the N.

Total Nitrogen-15 Recovery
The average recovery of winter-applied poultry manure 15N in soil after three growth seasons was 28% in the first experimental series and 42% in the second series (Table 3). As the crop recovery of winter-applied manure was almost the same in the two series (Fig. 6, Table 3), more 15N was totally recovered in crop and soil in the second series (62%) compared with the first series where 49% of manure 15N was recovered (Table 3). The differences in recovery between the two series were probably established in the first year of the two series. In the first series, precipitation was almost 40% higher from the day of application and until July compared with the second series. This may have promoted 15N leaching losses of the winter-applied poultry manure in the first series, but did not affect 15N recovery in the barley.

After application of poultry manure and mineral fertilizer in spring, soil recoveries of 15N from both N sources were considerably higher than after application of poultry manure in winter (Table 3). The recovery of 15N in soil amended with poultry manure in spring averaged 43 and 52% for the three manures in the first and second series, respectively, which resulted in a total 15N recovery in crop and soil after the three years of 94 to 95%. The average soil recovery of 15N applied in the two doses of mineral fertilizer was 26% in the first series and 45% in the second series, which gave a total recovery of mineral fertilizer in crops and soil of 84 and 92%, respectively (Table 3).

The bedding contained in two of the manures did not influence the recoveries and did not cause any measurable immobilization that could have reduced 15N losses. Losses are assumed mainly to have occurred by leaching in the first months after application rather than by gaseous losses via ammonia volatilization and denitrification. High amounts of ammonia may be lost if poultry manure is surface applied and not incorporated (Schilke-Gartley and Sims, 1993). The poultry manure in the present experiment was mixed with soil immediately. The high total N recoveries of spring-applied poultry manure N indicate that gaseous losses were low during this time of the year even though the higher temperatures in spring may have created favorable conditions for gaseous losses. The total recoveries of poultry manure 15N in crop and soil after the spring application are in accordance with Jensen et al. (1999), who likewise had a cropping system that included winter ryegrass. In contrast, Thomsen (2001) had considerably lower total soil and crop recoveries of 15N from solid manure 2 yr after application. The lower recoveries were probably a result of both the application time (September) and a crop sequence that included winter fallow between two cereals.

Shepherd and Bhogal (1998) concluded that autumn application of poultry manure should be avoided because of high leaching losses, whereas delaying the application until December diminished losses. Likewise, Chambers et al. (2000) stated that poultry manures should not be applied to free-draining soils in the period from autumn to early winter. The general low crop 15N uptake and 15N soil recovery of poultry manure applied in December or January in the present study shows that even this application time may result in high N losses. Although low temperatures slowed down mineralization, Sims (1986) found high proportions of mineralized N in poultry manure also at 0°C. Thus, the risk of N leaching losses is high when poultry manure is applied during cool and wet winters.


    CONCLUSIONS
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
For optimizing utilization of poultry manure, only spring application can be recommended whether or not the manure contains bedding with a high C:N ratio. If extensive N losses during storage have been avoided and the manure is applied in spring without significant gaseous N losses, a very high crop N utilization can be expected. With optimum conditions during storage and spreading, the potential N utilization is almost as high as can be obtained with mineral fertilizer.


    ACKNOWLEDGMENTS
 
This work was supported financially by the Ministry of Food, Agriculture and Fisheries (Project BÆR98-DJF-3).

Received for publication April 17, 2003.


    REFERENCES
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 




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Vadose Zone Journal
Journal of Plant Registrations Journal of
Environmental Quality
The Plant Genome