Agriculture, Ecosystems and Environment 99 (2003) 97–111

Nitrous oxide fluxes from maize fields: relationship to yield,

site-specific fertilization, and soil conditions
Ulrike Sehy∗ , Reiner Ruser, Jean Charles Munch
GSF-National Research Centre for Environment and Health, Institute of Soil Ecology,
Ingolstädter Landstrasse 1, D-85764 Neuherberg, Germany
Received 24 October 2001; received in revised form 19 March 2003; accepted 26 March 2003

Abstract
Nitrous oxide emissions and selected soil properties in a high and a low yielding area of a maize field were monitored
weekly over a 1-year period. In both the high and the low yielding area, N2 O emissions from a treatment subject to site-specific
N-fertilization were compared to a conventionally fertilized control.
Emission peaks were measured following N fertilization, rainfall, harvest, tillage and freeze-thaw cycles from all treatments
in conditions favorable for denitrification. Between 80 and 90% of annual emissions were released between April and Septem-
ber. A value of 60% WFPS was identified as a threshold for the induction of elevated N2 O emissions (>50 ␮g N2 O-N m−2 h−1 ).
A significant relationship (r 2 = 0.41) between N2 O flux rates and WFPS was found when neither soil nitrate contents nor
temperature were limiting for microbial denitrification.
Mean cumulative N2 O emissions from the control treatments in the high yielding area, located in a footslope position and
thus receiving lateral water and nutrient supply, more than doubled those from the control treatments in the low yielding
area in a shoulder position (8.7 and 3.9 kg N2 O-N ha−1 , respectively). Higher average WFPS in the high yielding area was
identified as responsible for this difference.
The site-specific fertilized treatments in the low yielding area were supplied with 125 kg N fertilizer ha−1 as compared to
150 kg N fertilizer ha−1 (control treatments). This reduction resulted in 34% less N2 O released in roughly 10 months following
differentiated fertilization while crop yield remained the same. In the high yielding area, N fertilizer supply in the site-specific
fertilized treatment was 175 kg N ha−1 as compared to 150 kg N ha−1 in the control. Neither crop yield nor N2 O emissions
were significantly affected by the different fertilizer rates.
© 2003 Elsevier Science B.V. All rights reserved.
Keywords: N2 O; Soil moisture; Soil nitrate; N fertilization; Site-specific fertilization; Precision farming

1. Introduction centration of 314 ppb and continues to increase (IPCC,

Owing to human activities, the concentration of the
greenhouse gas nitrous oxide (N2 O) in the troposphere
has increased from 270 ppb in 1750 to a current con-
∗ Corresponding author. Tel.: +49-89-544184-42;
fax: +49-89-544184-49.
E-mail address: sehy@oekom.de (U. Sehy).
2001b). In addition to its contribution to global warm-
ing (IPCC, 2001b), nitrous oxide is involved in the
destruction of the stratospheric ozone layer (Crutzen,
1981). Agriculture is estimated to contribute up to
80% to global anthropogenic N2 O emissions, more
than half of which is released from agricultural soils
(Mosier et al., 1998; Kroeze et al., 1999). The increase
in anthropogenic N2 O emissions is primarily the result

0167-8809/$ – see front matter © 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0167-8809(03)00139-7
98 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
of increased N input into agricultural soils (Mosier
et al., 1998; IFA and FAO, 2001). Between 1930 and
1990, the total anthropogenic N input through syn-
thetic and organic fertilizers, N2 -fixation, and the in-
put of plant residues rose from 44 to 180 Tg N a−1
(Kroeze et al., 1999).
Nitrification and denitrification, microbial pro-
cesses which rely on mineral N (ammonia and nitrate)
as their substrate, are responsible for most of the
N2 O produced in soil (Firestone and Davidson, 1989;
Hutchinson and Davidson, 1993; Bremner, 1997).
Consequently, following N-fertilizer application, an
increase in denitrification rates and/or N2 O flux rates
has been frequently found in field and laboratory
experiments (Ryden, 1983; Jambert et al., 1997;
Mulvaney et al., 1997; Smith et al., 1997; Kaiser et al.,
1998; Simojoki and Jaakkola, 2000). However, high
N2 O emissions often only occurred when soil water
content was high or immediately following rainfall
(Aulakh et al., 1984; Skiba et al., 1992; Hansen et al.,
1993; de Klein and van Logtestijn, 1994; Flessa et al.,
1995; Smith et al., 1998; Dobbie et al., 1999; Ruser
et al., 2001). Under these conditions, N2 O emissions
are mostly derived from microbial denitrification.
It has been stated that worldwide, agricultural emis-
sions can be most effectively reduced in high-intensity
agricultural systems by minimizing N-surpluses
(III) to assess the potential of site-specific fertilization
(IPCC, 2001a). This can be achieved through the
application of a range of measures aiming at synchro-
nizing N application with crop demand, including the
application of N-fertilizer taking into account soil-
and plant-N contents (Eichner, 1990; van Kessel et al.,
1993) and plant demand (Weier et al., 1993); the use
of intercrops (Wagner-Riddle and Thurtell, 1998); the
splitting of N-fertilizer input (Ruser et al., 2001) and
the use of more sophisticated fertilization techniques
(nitrification inhibitors, controlled release fertilizers)
(Mosier et al., 1996; Smith et al., 1997).
A comparatively new measure aimed at avoiding
N-surplus in agricultural soils is precision farming,
which, in practice, is mostly equivalent to site-specific
fertilization (Auernhammer, 2001). Common agricul-
tural practice in Germany is to apply N-fertilizer at
equal rates within often arbitrarily drawn field borders
irrespective of intrinsic heterogeneity. This may result
in zones that receive insufficient amounts of fertilizer
for maximum yield, and other areas that are oversup-
plied with agrochemicals, thus imposing a threat to
the environment (Maidl et al., 1999). Site-specific fer-
tilization, in contrast, takes into account that soil phys-
ical, chemical and biological heterogeneity within a
field may result in spatial and temporal variability in
nutrient cycling which is likely to have an impact on
crop yield as well as on gaseous N losses. With preci-
sion farming, variations in soil or crop characteristics
within a field are identified, mapped, and manage-
ment actions are spatially and temporally adjusted to
crop requirements in the respective areas (Dawson
and Johnston, 1997; Mandal and Ghosh, 2000).
To our knowledge, no investigations have been
published on the effects of site-specific fertilization
on N2 O emissions based on field measurements. We
present a study where, both in a high yielding area
and in a low yielding area of a field, N2 O emissions
from a uniformly fertilized treatment and from an
adjacent site specifically fertilized treatment were
compared. The aims of the study were:
(I) to identify the regulating parameters for N2 O
emissions from the investigated agricultural
soils;
(II) to quantify the N2 O emissions from a high and
a low yielding area of an arable field based
on year-round measurements depending on site
characteristics;
for reducing the emissions of nitrous oxide.
2. Materials and methods
2.1. Study site
The field experiment was carried out at the research
station Scheyern of the Munich Research Alliance on
Agroecosystems (FAM) between April 1999 and April
2000. The research station is located 445–498 m above
sea level in a hilly landscape derived from tertiary
materials and partly covered by loess, approximately
40 km north of Munich (48◦ N30.0 , 11◦ E20.7 ).
Study site was a gently undulating field cropped
to maize (Zea mays, L.) in 1999, followed by win-
ter wheat (Triticum aestivum, L.). For soil character-
istics refer to Table 1. Site-specific management on
this field was introduced in 1999. For that purpose five
classes of long-term stable crop yield were established
 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111 99

Table 1
Soil characteristics of the topsoil (0–20 cm) of the treatmentsa
High yielding Low yielding
area (HC, HP) area (LC, LP)
Soil classification (USDA Typic Dystric
taxonomy) Udifluvent Eutrochrept
pH (10−2 M CaCl2 )b 5.9 6.1
Corg (%)b 1.7 b 1.4 a
Nt (%)b 0.17 b 0.15 a
Grain size fractions (%)c
Clay < 2 ␮m 15 20
Silt 2–63 ␮m 49 51
Sand 63–2000 ␮m 36 29
a Different letters indicate significant differences between treat-

ments (t-test, P < 0.05).

b Own measurements.
c From Scheinost and Sinowski (1993).

(<85%, 85–95%, 96–105%, 106–115%, >115% of

average yield) from maps integrating yield during 3
years. The yield potentials of the three highest and the
three lowest yield classes were considered sufficiently
similar to justify a subsequent reduction to two areas
(J. Schächtl, pers. comm.)—a high yielding area con-
taining classes with >96% of average yield (66% of
total area) and a low yielding area containing classes
with ≤95% of average yield (34% of total area).
The field was then randomly subdivided into (a)
strips that were site-specifically fertilized (precision
farming (P)) and (b) strips that were uniformly fertil-
ized (conventional farming (C)) in order to permit a Fig. 1. Design of the field experiment. Dark gray: high yielding
comparison of the two fertilizer treatments (Fig. 1). area; white: low yielding area; P: strips fertilized according to
precision farming measures; C: strips fertilized according to con-
Strips were 7.5 m wide. The treatments investigated ventional measures; HP, HC, LP, LC: location of the treatments;
were high yielding area, uniformly fertilized (HC); circles: location of gas flux measurement chambers.
high yielding area, site specifically fertilized (HP); low
yielding area, uniformly fertilized (LC); and low yield-
ing area, site specifically fertilized (LP) (Fig. 1). The potential (derived from yield maps) and the result-
two treatments in the high yielding area were situated ing average plant demand. The calculated N-fertilizer
in a footslope position, corresponding to the highest amount of 150 kg N ha−1 corresponded to the aver-
yield class (>115% of average yield), whereas the two age plant demand minus the average N-supply from
sites in the low yielding area were on a rounded hill- the soil (derived from grid sampling) (Scheinost and
top, corresponding to the second lowest yield class Sinowski, 1993). In the following, the uniformly fer-
(85–95% of average yield). We considered the lowest tilized strips (C) are referred to as the control. For the
yield class not to be representative of the low yielding site specifically fertilized strips (P), the amounts of
area, since it had included only a small proportion of N-fertilizer were calculated so that in the high yielding
the total field that corresponded exclusively to areas area, maximum yield potential was expected to be 20%
close to the field border. higher than field average. To satisfy the corresponding
For the uniformly fertilized strips (C), the amount higher crop N demand, 20% more N-fertilizer should
of N-fertilizer was determined from the average yield be applied (180 instead of 150 kg N ha−1 ). However,
100 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111

due to lateral supply and higher organic N-content Table 2

(Table 1), the soil N-supply in the high yielding area Management of the fielda
is assumed to be 5 kg N ha−1 higher than field aver- 8 August 1998 Harvest of winter wheat
age. Therefore, 175 kg N ha−1 were applied to the P 14 August 1998 Sowing of mustard (intercrop)
strips in the high yielding area. In the low yielding 4 May 1999 Sowing of maize
area, yield potential was expected to be 20% lower Fertilizer AP (banded)
than field average, resulting in 20% less N-fertilizer 40 kg N ha−1 (all treatments)
to be applied (120 instead of 150 kg N ha−1 ). The ac- 1 June 1999 Fertilizer CAN (broadcast)
HC: 110 kg N ha−1
tual amount of N-fertilizer applied was increased by a HP: 135 kg N ha−1
value of 5 kg N ha−1 (to 125 kg N ha−1 ), because soil LC: 110 kg N ha−1
N-supply in the low yielding area was expected to be LP: 85 kg N ha−1
5 kg N ha−1 lower than field average. 19 September 1999 Harvest of maize
22 October 1999 Tillage (0–5 cm) and
The investigation period was examined in total as
sowing of winter wheat
well as subdivided into four periods as follows:
24 March 2000 Fertilizer ANU (broadcast)
A. complete investigation period (23 April 1999–24 HC: 50 kg N ha−1
April 2000); HP: 35 kg N ha−1
B. before differentiated fertilization of maize (23 LC: 50 kg N ha−1
LP: 50 kg N ha−1
April–31 May 1999);
a CAN: calcium-ammonium-nitrate; AP: ammonium-phos-
C. from differentiated fertilization of maize to harvest
of maize (1 June–18 September 1999); phate; ANU: ammonium-nitrate-urea solution (25/25/50% N).
D. from harvest of maize to first fertilization of winter rows (data not shown). Gas sampling was performed
wheat (19 September 1999–23 March 2000); at least once a week beginning at 10 a.m. on each
E. from first fertilization of winter wheat to end of sampling day. During the sampling period, while the
investigation period (24 March–24 April 2000). base rings were covered with dark closed chambers
(height 10–15 cm) for 60–90 min, we withdrew four
2.2. Determination of N2 O fluxes 100 ml samples into evacuated glass flasks. The first
sample was taken immediately after chamber closure,
For each treatment (HC, HP, LC, LP) five polyvinyl subsequent samples at equal intervals (20–30 min de-
chloride base rings (height 15 cm; inner diameter pending on sampling period) during chamber closure.
30 cm) were permanently installed in the soil down For a more detailed description of chamber design and
to a depth of 10 cm and distributed across an area sampling scheme refer to Flessa et al. (1995). Sam-
of 7.5 m × 15 m (Fig. 1). Each ring enclosed a sur- ples were analyzed for CO2 and N2 O concentrations
face area of 706.5 cm2 . The base rings were installed using an automated gas chromatographic system with
between two rows of maize plants or enclosing the a 63 Ni electron capture detector (ECD) as described
wheat plants. The rings were removed immediately in detail by Loftfield et al. (1997). Gas fluxes were
prior to soil management operations, and immediately calculated as rates of observed linear concentration
re-installed afterwards in the same location. Gas fluxes change per time and surface area and expressed as
were measured at least 6 days after the base rings were arithmetic means (n = 5) with standard deviation.
installed so that the impact of the installation would
have minimal impact on the gas fluxes. Since the first, 2.3. Climate and soil variables
uniform N application was banded to the seed row of
maize (Table 2), measurement of N2 O fluxes between Soil temperature was measured at the same time
two seed rows might have led to an underestimation as gas sampling at 10 cm depth in each treatment.
of total flux. Therefore, additional measurements in Air temperature and precipitation data were supplied
the seed row were made, which showed that annual by the meteorological station of the research station
N2 O flux might be underestimated by up to 28% (TU Munich, Chair of Bioclimatology and Immission
if measurements are restricted to between the seed Research).
 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
During the gas sampling period, we took bulk soil
samples from 0 to 10 cm depth by combining 5–10 soil
cores. Samples were kept cool for transportation and
processed on the same day. Soil moisture was deter-
mined gravimetrically after drying for 24 h at 105 ◦ C.
Water-filled pore space (WFPS) was calculated using
the measured soil bulk density data (arithmetic means
of four measurement dates) and assuming a particle
density of 2.65 g cm−3 .
Fresh soil was extracted with 10−2 M CaCl2 (soil to
solution ratio 1:2 (w/v)), shaken in a rotary shaker for
1 h, centrifuged at 4000 rpm for 10 min, and filtered
through a 0.45 ␮m membrane filter. Extract solution
was split and immediately analyzed for dissolved or-
ganic carbon (DOC) in a TOC analyzer (Shimadzu
TOC-5050A) or stored frozen until analyzed for NH4 +
and NO3 − in a continuous flow analyzer (Skalar An-
alytical SA 20/40).
2.4. Statistical analysis
Relationships between variables were assessed us-
ing stepwise linear regression analysis (P < 0.05).
Normal distributions of flux rates and driving vari-
ables were tested using the Kolmogorov–Smirnov
test at P > 0.05. Variables such as N2 O flux rates,
CO2 flux rates and soil nitrate contents were log nor-
mally distributed and were log10 -transformed before
statistical analysis. Since we used dark chambers for
gas accumulation, CO2 flux rates were only included
for statistical analysis for periods where no plants
were present in the soil covers (April—November
1999).
Cumulative fluxes were calculated for each of the
five flux chambers per treatment assuming measured
fluxes to be steady between two sampling dates. To
assess significant differences between treatments, cu-
mulative fluxes were subjected to pairwise multiple
comparisons using the Student–Newman–Keuls test at
a significance level P < 0.05.
For each treatment (HP, HC, LP, LC) we calculated
the quotient of cumulated N2 O emissions (periods C
and D) and maize yield as an environmental indicator.
We obtained the quotients by randomly dividing cu-
mulative N2 O emissions from individual soil covers
(n = 5) by yield data from individual plots (n = 5) and
compared them using the Student–Newman–Keuls test
at a significance level P < 0.05.
101
3. Results and discussion
3.1. Temporal changes in N2 O flux rates
depending on regulating parameters
Average air temperature in the investigated year was
8.8 ◦ C, the sum of precipitation amounted to 937 mm.
Both parameters surpassed the long-time averages—
7.4 ◦ C for temperature and 833 mm for precipitation—
measured at the research station.
The temporal course of N2 O emissions from the
four treatments (Figs. 2 and 3) is characterized by
low flux rates (<50 ␮g N2 O-N m−2 h−1 ) for most
sampling dates, interrupted by several high emis-
sion periods. Differing in magnitude and duration,
these distinct periods were observed both in the
high and the low yielding area, indicating common
controllers. Maximum emission rates were around
1400 ␮g N2 O-N m−2 h−1 in the high and around
1300 ␮g N2 O-N m−2 h−1 in the low yielding area.
Generally, highest flux rates occurred in spring and
summer following N fertilization and were associated
with rainfall. After the first N application in May 1999,
elevated N2 O flux rates occurred after 25 mm rain had
fallen within 6 days. Fluxes were very high from the
high yielding area (up to 1400 ␮g N2 O-N m−2 h−1 )
and lower and less persistent from the low yielding
area (up to 250 ␮g N2 O-N m−2 h−1 ) (Figs. 2 and 3).
During 2 weeks following the second N-fertilization
in early June, N2 O flux rates were only slightly el-
evated in both areas. These relatively low emission
rates following N input can be attributed to low soil
water contents, which fell below 50% WFPS during
that period. According to Davidson (1991), the domi-
nant process for N2 O evolution under these conditions
is nitrification. High N2 O flux rates, in contrast, are
usually associated with the process of denitrification
(Firestone and Davidson, 1989), which becomes dom-
inant for N2 O production at WFPS values of 60%
(Davidson, 1991). Strongly elevated flux rates in both
areas did not take place until heavy rain in mid-July
(66 mm within 4 days) resulted in WFPS values above
60% in all treatments (Figs. 2 and 3). Cumulative
N2 O emissions within 2 weeks following the onset of
rain amounted to 30% (high yielding area) and 60%
(low yielding area) of total annual N2 O emissions.
This underlines the importance of rainfall for N2 O
emissions, which has been frequently described for
102 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111

Fig. 2. N2 O flux rates, CO2 flux rates, soil nitrate contents, soil WFPS and daily precipitation of the high yielding area treatments. Arrows
with numbers indicate fertilizer amounts (kg N ha−1 ) applied to the respective areas.

agricultural soils (Cates and Keeney, 1987; Ball et al.,
1999; Dobbie et al., 1999; Baggs et al., 2000; Scott
et al., 2000; Ruser et al., 2001). In our study, 85% of
all elevated flux rates (>50 ␮g N2 O-N m−2 h−1 ) were
measured at soil water contents between 55 and 90%
WFPS, with maximum emissions at 65% WFPS (cf.
Fig. 4A). Similarly, Simojoki and Jaakkola (2000)
measured highest N2 O emissions at WFPS between
60 and 90% in agricultural soils and Dobbie et al.
(1999) between 70 and 90% in agricultural soils and
fertilized grassland. In the present study, flux rates
above 50 ␮g N2 O-N m−2 h−1 and at WFPS below
60% were only measured when WFPS was above this
value at the preceding measurement date. Under these
conditions, inner aggregates are still waterlogged and
sustain denitrification, even if a large share of pores
between aggregates is already air-filled (Smith, 1980).
We thus consider 60% WFPS the threshold value for
the induction of elevated N2 O flux rates through den-
itrification in our soils, a value somewhat lower than
the threshold of 65% WFPS found by Clayton et al.
(1997). High CO2 fluxes were measured during the pe-
riod of highest flux rates in July and indicate substan-
tial heterotrophic activity after rewetting of the dry soil
as a result of increased carbon availability (Sexstone
et al., 1988; Rudaz et al., 1991). Heterotrophic activity
 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111 103

Fig. 3. N2 O flux rates, CO2 flux rates, soil nitrate content, soil WFPS and daily precipitation of the low yielding area treatments. Arrows
with numbers indicate fertilizer amounts (kg N ha−1 ) applied to the respective areas.

enhances O2 consumption and promotes anaerobic
conditions; additional high soil nitrate and water
contents favored high N2 O losses through denitrifi-
cation in that period (Firestone and Davidson, 1989;
Hutchinson and Davidson, 1993; Granli and Bøckman,
1994).
Fig. 4A shows that WFPS and N2 O emissions were
not related when all data were considered. From a
scatter plot of nitrate content versus N2 O emissions,
we identified a nitrate content of 5 kg NO3 − -N ha−1
(about 3.5 mg NO3 − -N kg−1 d.wt.) as the lower
threshold for the induction of elevated N2 O emissions
above 50 ␮g N2 O-N m−2 h−1 (data not shown). Us-
ing only N2 O flux rates from dates where soil nitrate
contents were above this value, a small, but signifi-
cant relationship between N2 O flux rates and WFPS
was found (r2 = 0.13) (Fig. 4B). This relationship
improved to r 2 = 0.41 when additionally all data
were excluded where soil temperature was below
7 ◦ C (Fig. 4C), reflecting that microbial nitrification
and denitrification are usually limited at low soil tem-
peratures (Granli and Bøckman, 1994). The findings
presented here are in accordance with other studies,
where no relationship between N2 O emissions and
WFPS was found when parameters like soil mineral
N and/or C content or soil temperature were limiting
104 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
Fig. 4. Scatterplots of N2 O flux rates versus WFPS. Horizontal
line denotes N2 O flux rates >50 ␮g N2 O-N m−2 h−1 , vertical lines
delimit WFPS values between 55 and 90%. (A) All data (n = 233),
(B) dates where soil nitrate contents were >5 kg N ha−1 , (C) dates
where soil nitrate contents were >5 kg N ha−1 and soil temperature
was >7 ◦ C.
N2 O production (Ryden, 1983; Clayton et al., 1997;
Dobbie et al., 1999; Conen et al., 2000). The thresh-
old of 3.5 mg NO3 − -N kg−1 d.wt. found in this study
is somewhat lower than the values reported in the
literature. In the studies by Ryden (1983) and Dobbie
et al. (1999), denitrification and/or N2 O flux rates
tended to be small below 5 mg NO3 − -N kg−1 d.wt.,
however, no exact values for maximum N2 O flux
rates were given. Conen et al. (2000) reported a value
of 10 mg NO3 − -N kg−1 d.wt. to limit N2 O flux rates
to below 42 ␮g N2 O-N m−2 h−1 .
We applied a stepwise multiple linear regression to
analyze the relationships between N2 O emissions and
the parameters soil nitrate content, DOC content, CO2
flux, temperature, WFPS and the cumulated amount of
precipitation within 1 week before each gas sampling
(PPcum ) (Table 3). In periods B and C—during vege-
tative growth—CO2 flux and PPcum explained about
70% of the N2 O flux variability in the high yield-
ing area and in treatment LC in the low yielding area
(Table 3). WFPS and temperature explained 83% of
the N2 O flux variability of treatment LP. Supporting
the observed high influence of rain for the induction
of flux peaks, PPcum generally had more influence on
N2 O flux than WFPS. In contrast to WFPS, the pa-
rameter PPcum takes into account that on the one hand
N2 O emissions may be low at high soil water status,
because nitrification is limited and/or denitrification
proceeds rapidly to N2 (Sahrawat and Keeney, 1986)
and on the other hand that flux peaks are frequently in-
duced by an abrupt change in soil water status whether
or not soil water content is already above the threshold
level (Mosier et al., 1991; Clayton et al., 1994).
After the fertilizer-induced emission peaks in
spring and summer, elevated N2 O flux rates were
again measured in September after harvest. Preceding
the emission peaks, the soil had dried to the lowest
water content during the investigated period and was
well below 50% WFPS in all treatments (Figs. 2
and 3). High emissions were induced upon rewetting
of the soil to values of 65% (low yielding area) and
75% WFPS (high yielding area). At the end of Oc-
tober, after shallow tillage, sowing of winter wheat
and with air temperatures rising to >10 ◦ C, high CO2
emissions were accompanied by a distinct rise in
N2 O emissions in both areas (Figs. 2 and 3). This can
be explained by the fact that tillage increases aera-
tion and physically disturbs soil aggregates and crop
residues, thus rendering additional substrates avail-
able for microbial attack and promoting microbial
activity (Granli and Bøckman, 1994; Kristensen et al.,
2000). Emissions peaked after WFPS surpassed 60%,
again indicating an increasing share of denitrification
in N2 O production (Davidson, 1991).
By the end of January 2000, average daily tem-
peratures rose from −14 ◦ C to +8 ◦ C. In the week
before, the soil had remained slightly frozen to a
 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111 105

Table 3
Coefficients of determination (in brackets: r2 ) for stepwise multiple linear regressions of N2 O flux rates versus soil parameters differentiated
according to the period of vegetative growth (periods B + C) and winter period (period D) (periods as defined in Section 2.1)a
Period HP HC LP LC

B + C (n = 23) CO2 (0.60) CO2 (0.60) WFPS (0.65) PPcum b (0.55)

PPcum (0.73) PPcum (0.69) T (0.83) CO2 (0.72)
NO3 (0.86)
D (n = 25) n.s.c T (0.36) T (0.43) T (0.32)
NO3 (0.53) PPcum (0.55) PPcum (0.46)
a Displayed are parameters that are included in the linear model at a F-to-enter-probability < 0.05 and F-to-remove probability > 0.1,

numbers before parameters indicate the order of parameters as entered in the model.
b Cumulated amount of precipitation within 1 week before each gas sampling.
c Not significant.

depth of 10 cm. The thawing of a 20 cm snow cover
as well as 14 mm rain on 30 January resulted in
complete thawing of the soils and in very high soil
water contents in all treatments. On 31 January, el-
evated N2 O flux rates were measured from all treat-
ments. Emissions were three to six times higher than
the background flux measured in the 2 months be-
fore. They amounted to 120 ␮g N2 O-N m−2 h−1 and
were comparable to values reported by Cates and
Keeney (1987) for fertilized maize fields [150 ␮g
N2 O-N m−2 h−1 (only soil surfaces thawed) and
200 ␮g N2 O-N m−2 h−1 (soils completely thawed)].
However, the freeze–thaw related flux rates reported
in this study did not reach the magnitude previ-
ously described from winter measurements at the
same experimental station: Following freeze–thaw
events, Ruser et al. (1998, 2001) measured 500 and
>900 ␮g N2 O-N m−2 ha−1 , respectively, and Flessa
et al. (1995) found >2000 ␮g N2 O-N m−2 ha−1 . We
attribute the relatively low N2 O emissions found in
our study to the complete water logging during that
period, since under conditions of very high water
status gas exchange between soil and atmosphere is
hindered and the complete reduction of N2 O to N2 is
promoted (Granli and Bøckman, 1994). In the winter
period D, N2 O emissions were significantly related to
soil temperature in three of the four treatments (HC,
LP, LC) (Table 3). This relates to the observation that
N2 O emissions were generally low during winter, and
that high flux rates in period D were associated with
either: (a) rising temperatures after tillage, leading to
enhanced substrate availability (at the end of Octo-
ber) or (b) rising temperatures during thawing of the
frozen soil.
3.2. Cumulative N2 O emissions depending
on site characteristics
In the complete investigation period A, signifi-
cantly more N2 O was released from the treatments
in the high yielding area than from those in the low
yielding area (Table 4). Although treatments HC and
LC received the same amount of fertilizer, cumulative
N2 O emissions from HC more than doubled those
from LC (8.7 and 3.9 kg N2 O-N ha−1 ). One reason
for the different emission levels between high and
low yielding area sites may be the persistently higher
soil water content in the former (Figs. 2 and 3). Since
the soil textures of both high and low yielding areas
are not strikingly different (Table 1), this parameter is
not likely responsible for the difference in soil water
content. We ascribe the higher soil water content in
the high yielding area to the location in a footslope
position, which results in lateral water supply and
temporary reduced percolation. Corre et al. (1996) and
Pennock and Corre (2001) also suggest that the higher
WFPS in footslope positions is responsible for higher
N2 O evolution as compared to shoulder positions.
In their investigations, different emission levels were
constantly associated with certain landscape positions
and topography was considered an indirect driving
variable, influencing the processes and conditions that
control microbial denitrification rates and N2 O emis-
sions (i.e. water and nutrient availability) (Pennock
et al., 1992; van Kessel et al., 1993; Corre et al., 1996).
In our study, a value of 60% WFPS was iden-
tified as non-limiting for the induction of elevated
N2 O emissions (>50 ␮g N2 O-N m−2 h−1 ). The key to
higher N2 O emissions from the high yielding area may
 106
U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
Table 4
Cumulative N2 O emissions from the treatmentsa
Period HP HC LP LC

N2 O emissions Fertilizer N2 O emissions Fertilizer N2 O emissions Fertilizer N2 O emissions Fertilizer

(kg N2 O-N ha−1 ) (kg N ha−1 ) (kg N2 O-N ha−1 ) (kg N ha−1 ) (kg N2 O-N ha−1 ) (kg N ha−1 ) (kg N2 O-N ha−1 ) (kg N ha−1 )
A 10.1 b ± 2.0 210 8.7 b ± 1.2 200 3.1 a ± 0.6 175 3.9 a ± 0.5 200
B 3.4 c ± 1.1 (38.7) 40 2.4 b ± 0.7 (30.0) 40 0.5 a ± 0.2 (19.3) 40 0.3 a ± 0.1 (7.2) 40
C 4.2 c ± 0.8 (47.2) 135 3.8 c ± 0.4 (48.7) 110 2.0 a ± 0.2 (70.4) 85 3.0 b ± 0.5 (81.4) 110
D 1.3 b ± 0.4 (14.2) – 1.7 b ± 0.5 (21.3) – 0.3 a ± 0.2 (10.4) – 0.4 a ± 0.1 (11.4) –
E 1.3 b ± 0.5 35 0.8 b ± 0.5 50 0.3 a ± 0. 50 0.2 a ± 0.1 50
a Data are shown as arithmetic means with standard deviation; data in parentheses: percent of maize-related emission (periods B and D: 23 April 1999–23 March 2000);

different letters indicate significant differences between treatments.

 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
Fig. 5. WFPS values of all treatments during periods B and
C. Displayed are 25 and 75%-percentiles as vertical boxes, 10
and 90%-percentiles as whiskers and outliers as circles. Within
individual boxes, the median is displayed as solid line and the
arithmetic means as dotted line. The solid line across the figure
represents 60% WFPS, a value which was identified as non-limiting
for the induction of elevated N2 O emissions.
be that during periods B and C WFPS was above this
value on the majority of measurement dates (Fig. 5),
whereas in the low yielding area, the majority of ob-
servations showed WPFS values below 60%, indicat-
ing that for most of the dates N2 O production from
denitrification was limited by low WFPS.
Apart from soil water contents, treatments HC
and LC differed in soil nitrate content. This was
the case despite the same amounts of N applied in
both treatments and probably is the result of higher
organic N contents (Table 1) as well as lateral N
supply from the surrounding slopes in the high yield-
ing area. However, we assume that the different soil
nitrate contents in the high and low yielding area
were not responsible for different N2 O emissions,
since soil nitrate did not explain much of N2 O flux
variability in periods B and C (Table 3), although the
results of the stepwise multiple regression analysis—
strong positive relationships of N2 O flux rates with
CO2 flux rates and soil water contents—point to
Table 5
Cumulative N2 O emissions (periods C and D), amount of N applied, maize yield and N2 O loss per yielda
Treatment
HP
N2 O emissions (kg N2 O-N ha−1 ) 5.4 c ± 0.7
N-fertilizer (kg N ha−1 ) 175
Yield (t d.wt. ha−1 ) 20.9 a ± 2.1
N2 O per yield (kg N2 O-N t−1 d.wt.) 0.26 c ± 0.04
a Data are shown as arithmetic means with standard deviation (n = 5); different letters indicate significant differences between treatments.
107
microbial denitrification as the dominant source for
high N2 O emissions. In the literature, values of
10 mg NO3 − -N kg−1 d.wt. (Mosier et al., 1983) and
25 mg NO3 − -N kg−1 d.wt. (Limmer and Steele, 1982)
are given as thresholds above which denitrifica-
tion rates are independent of soil nitrate contents.
In our study, average soil nitrate contents in peri-
ods B and C were 48 (HP), 36 (HC), 22 (LP) and
27 (LC) mg NO3 − -N kg−1 d.wt. and thus above the
threshold value of 25 mg NO3 − -N kg−1 d.wt. except
for treatment LP. This suggests that nitrate contents in
treatments HP, HC and LC were mostly non-limiting
for microbial N2 O production. This is corroborated
by the fact that only in treatment LP was soil nitrate
content included in the multiple regression analysis.
Supporting these results, the largest difference in N2 O
emissions between HC and LC occurred between
May and the end of July, while at the same time N2 O
emissions were the same in HC and HP, although the
difference in soil nitrate contents between HC and HP
was much larger than between HC and LC (Figs. 2
and 3).
3.3. The influence of site-specific fertilization on
cumulative N2 O emissions and maize yield
In the high yielding area, during the approximately
10 months following N application (periods C and
D), cumulative N2 O emissions were not different
between HP and HC (Table 5). Average N2 O flux
rates during that period were 80 ␮g N2 O-N m−2 h−1
from both treatments. In contrast, in the low
yielding area, 34% less N2 O was emitted from
treatment LP (2.3 kg N2 O-N ha−1 ) than from LC
(3.5 kg N2 O-N ha−1 ). Beginning with differentiated
fertilization, flux rates from LC surpassed those from
LP on nearly every measurement date until flux rates
HC LP LC
5.5 c ± 0.5 2.3 a ± 0.3 3.5 b ± 0.6
150 125 150
19.3 a ± 2.6 18.2 a ± 1.6 17.7 a ± 1.5
0.28 c ± 0.02 0.12 a ± 0.02 0.20 b ± 0.03
108 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
in both treatments decreased to background levels
(<5 ␮g N2 O-N m−2 h−1 ) in November.
In treatment HP, the amount of N-fertilizer was in-
creased by 25 kg N ha−1 as compared to the conven-
tionally fertilized control HC. In treatment LP, the
amount of N-fertilizer was reduced by 25 kg N ha−1
as compared to the control LC (Table 5). Accordingly,
average soil nitrate contents were higher in HP than
in HC (48 versus 36 kg N ha−1 in period C) and lower
in LP than in LC (22 versus 27 kg N ha−1 ), especially
in the period of vegetative growth.
Above, we have ascribed the lack of correlation be-
tween soil nitrate contents and N2 O emissions to the
fact that soil nitrate contents were above threshold
values limiting microbial denitrification (Limmer and
Steele, 1982). We suggest that this is also the reason
for N2 O emissions from both high yielding area treat-
ments being the same despite different fertilizer rates
and nitrate contents. Only in the low yielding area,
where N2 O emissions from treatment LP were a func-
tion of soil nitrate contents (Table 3), we calculated a
significant difference between conventional and pre-
cision farming treatment.
Maize yield was not affected by the three fertil-
izer rates (125, 150, 175 kg N ha−1 ) (Table 5). This
shows that soil N contents were not limiting crop
yield. Similar results were reported by Chantigny et al.
(1995), where an increase in N amounts from 120 to
180 kg N ha−1 hardly affected maize yield (no differ-
ence in a sandy clay; a slight increase of 5–8% in a
sandy loam). In a study by Isfan et al. (1995) the opti-
mum N-fertilizer rate for maize was between 115 and
146 kg N ha−1 , which is lower than the amounts used
in the C-treatments in this study. According to Liang
and MacKenzie (1994), the N use efficiency of maize
decreases with increasing amounts of N-fertilizer. This
might explain why the difference in average soil nitrate
contents between C and P treatments was 10 kg N ha−1
in the high yielding area and only 5 kg N ha−1 in the
low yielding area.
We related cumulated N2 O losses (periods C and D)
to maize yield (dry matter at harvest), in order to assess
the environmental impact of the different treatments
in relation to economic returns. Comparing high and
low yielding areas, the ratio of N2 O:yield was signifi-
cantly higher in the HC treatment (0.28 kg N2 O-N t−1 )
as compared to the LC treatment (0.20 kg N2 O-N t−1 )
(Table 5). These results have to be assessed taking
into account the climatic conditions in the investigated
year, which were characterized by higher temperatures
and especially more precipitation than average. In our
study year, maize yields were the same in both areas,
although the yield areas had been previously identi-
fied based on long-term differences in crop yield. In
a drier year, location in a shoulder position (= low
yielding area) is likely to result in water stress and
lower crop yields. On the other hand, in a drier year,
N2 O emissions from the high yielding areas are likely
to be lower, because soil moisture content will more
frequently limit high N2 O emissions through denitrifi-
cation and thus the N2 O:yield difference between the
two areas is likely to be smaller. Comparing the C and
P treatments, our data show that in the low yielding
area the ratio of N2 O:yield was significantly reduced
in the LP treatment as compared to LC (Table 5), while
in the high yielding area, this ratio was not affected by
differentiated fertilization. We assume that these rela-
tionships were not affected by the climatic conditions
that deviated from average in the study year.
In summary, in the low yielding area N surpluses
could be avoided through the adoption of site-specific
fertilization, resulting in significantly reduced N2 O
emissions and constant crop yield. In the high yield-
ing area, our data suggest soil N contents in both
treatments were on average above threshold levels
for both maximum crop yield and maximum N2 O
emissions.
The share of the low yielding area of the total field
is 34% (2.04 ha). Based on the results obtained from
the low yielding area treatments, a reduction of fertil-
izer amounts by 25 kg N ha−1 in the low yielding area
can result in a reduction of N2 O emissions by—in
total—2.44 kg N2 O-N within 10 months after differ-
entiated fertilization. Assuming unchanged N2 O emis-
sions from the high yielding area, the reduction of
N2 O evolved from the low yielding area corresponds
to a reduction by 0.41 kg N2 O-N ha−1 based on the
total field.
4. Conclusions
Higher soil water contents in the high yielding
area of a maize field resulted in significantly higher
N2 O emissions than in the low yielding area. The
location of the high yielding area treatments in a
 U. Sehy et al. / Agriculture, Ecosystems and Environment 99 (2003) 97–111
footslope position was responsible for the persis-
tently higher WFPS, pointing to the importance of
the landscape position for the quantification of N2 O
emissions.
Through the application of site-specific fertiliza-
tion, N input was increased by 17% in the high yield-
ing area and reduced by 17% in the low yielding area
as compared to the control. In the high yielding area,
neither crop yield nor N2 O emissions were affected
by higher N input, presumably because soil nitrate
contents were above levels limiting crop yield or
N2 O production. However, in the low yielding area,
the lower precision based N input in LP as compared
to LC resulted in 34% less N2 O released in roughly
10 months following differentiated fertilization,
while crop yield was not affected. We conclude that
site-specific fertilization has the potential to reduce
N2 O emissions. However, our results clearly demon-
strate that besides soil nitrate contents, lateral nutrient
and water supply and annual variations in climate—
especially rainfall patterns—have to be taken into
account for spatially differentiated N-fertilizer
recommendations.
Maize yields were the same in both areas, although
the areas had been previously differentiated based on
long-term differences in crop yield. We attribute this to
a relatively wet year, where water supply was sufficient
and location in a shoulder position (= low yielding
area) did not result in water stress and yield depression.
Acknowledgements
The scientific activities of the FAM Research Al-
liance on Agroecosystems are financially supported by
the German Federal Ministry of Education, Science,
Research and Technology (BMBF). Rent and oper-
ating expenses of the research station Scheyern are
paid by the Bavarian State Ministry for Education and
Culture, Science and Art. We thank Axel Berkenkamp
and Karl Zeller for their help in the field and in the
laboratory, Rolf Schilling and Brigitte Saueressig for
skilled technical assistance, Martin Winterhalter for
climate data as well as Josef Schächtl and Franz-Xaver
Maidl for yield data. We also thank Heiner Flessa,
Jens Dyckmans and two anonymous reviewers for
valuable comments on an earlier version of the
manuscript.
109
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