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Retention of deposited NH 4‐N and NO 3‐N in
coniferous forest ecosystems in Southern Sweden

S. Ingvar Nilsson , Dan Berggren & Olle Westling

To cite this article: S. Ingvar Nilsson , Dan Berggren & Olle Westling (1998) Retention
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of deposited NH 4‐N and NO 3‐N in coniferous forest ecosystems in Southern Sweden,
Scandinavian Journal of Forest Research, 13:1-4, 393-401, DOI: 10.1080/02827589809382999

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Scand. J. For. Res. 13: 393-401, 1998

Retention of Deposited NH+4-N and NO-3-N in Coniferous Forest

Ecosystems in Southern Sweden
S. INGVAR NILSSON1, DAN BERGGREN1 and OLLE WESTLING2
1
Department of Soil Sciences, Swedish University of Agricultural Sciences, P.O. Box 7014, SE-75007 Uppsulu, Sweden and
2
Swedish Environmental Research Institute, Aneboda Research Station, SE-360 30 Lammhult, Sweden

Scandinavian Journal
of Forest Research
Nîlsson, S. I.1, Berggren, D.1 and Westling, O. 2 ( 1 Department of Soil Sciences, Swedish
University of Agricultural Sciences, P.O. Box 7014, SE-75007 Uppsala, Sweden and 2Swedish
Environmental Research Institute, Aneboda Research Station, SE-360 30 Lammhult, Sweden).
Retention of deposited NH+4-N and NO-3-N in coniferous forest ecosystems in southern Sweden.
Received Feb. 4, 1997. Accepted Apr. 26, 1998. Scand. J. For. Res. 13: 393-401, 1998.
The main aim of the study was to investigate the retention pattern of atmospherically
deposited NH+4 plus NOf in podsolized forest soils in southern Sweden during the period
1985-1994. Nitrogen budgets were calculated for 37 coniferous forest sites dominated by
Norway spruce (Picea abies (L.) Karst.) or Scots pine (Pinus sylvestris L.). Total deposition of
NH+4 plus N O f was obtained by adding bulk deposition and dry deposition. Dry deposition
was calculated by using regional data on air concentrations of paniculate and gaseous N
compounds and previously established deposition velocities in stands of Norway spruce and
Scots pine. Nitrogen leaching was obtained by combining measured NH+4 and NO-3 concen-
trations with the amount of percolating soil water. The latter was chiefly estimated from the
Cl- balance, assuming that the flux in throughfall was equal to the C\~ flux in the soil water.
Nitrogen uptake by the trees was estimated by inserting data on diameter growth in empirical
biornass functions and then applying typical N concentration values gained from the literature.
The average N deposition was 13.3 ± 7.4 kg N h a - 1 yr-1 (±SD). Nitrogen leaching,
estimated annual soil N retention (maximum N immobilization), ecosystem N retention and
bole N accumulation (N uptake in bole wood + bole bark) were positively correlated with N
deposition. Bole N accumulation and ecosystem N retention were also negatively correlated
with stand age. The seven stands that showed a measurable amount of N leaching had a total
N deposition of > 15 kg N h a - 1 y r - 1 and were all situated in the southernmost part of the
investigated area. The average value of the estimated soil N retention. 8.5 ± 6.0 kg N ha-1
yr-1 was close to the range of default values used in estimates of critical N loads to European
forest ecosystems.
Key words: coniferous forest ecosystems, critical loads, default values, N
deposition, N immobilization, N leaching, N saturation. Picea abies, Pinus sylvestris.

INTRODUCTION the latter authors recently developed a model called

The long-term capacity of a forest ecosystem to cap-
ture N deposited as NH4+ or NOf is one of the main
factors to be considered in assessments of critical
loads of N, regardless of whether the critical load
refers to soil acidification or to soil eutrophication
(cf. Downing et al. 1993, Posch et al. 1995). One key
concept is N saturation, the definition of which has
been a matter of debate (Miller & Miller 1988).
Probably the best definition, which will be used in
this paper, was proposed by Ágren & Bosatta (1988).
They defined N saturation as the stage where long-
term N leaching is equal to or exceeds N deposition.
Using a similar approach, Aber et al. (1998) defined a
N cycling steady state with N leaching equal to N
deposition and with N uptake by the vegetation equal
to net N mineralization. Based on previous experi-
mental and theoretical work (e.g. Aber et al. 1991)
PnET-CN (Aber et al. 1997) which can be used to
simulate the time course for obtaining this steady
state under different N-deposition and land-use his-
tory scenarios.
The ideal tool for predicting critical loads of N or
the N-saturation level would be a fully mechanistic
N-cycling model with N deposition as one of the
driving variables. Examples of such modelling ap-
proaches are Soil-N (Eckersten et al. 1995) and the
PnET-CN model mentioned above. However, our
mechanistic understanding of some of the key pro-
cesses in the N cycle, such as N immobilization and
nitrification, will have to be improved before these
models can become fully operative. A combination of
experimental process studies and compilations of
data collected from comprehensive ecosystem investi-
gations is probably necessary in order to identify

1998 Scandinavian University Press. ISSN 0282-7581

394 S. I. Nilsson et al.
general relationships that could be useful for predict-
ing N-saturation levels and critical loads of N.
Johnson (1992) made a compilation of N cycling
investigations in the northern temperate and boreal
zones. Most of the studies were carried out either in
the USA or in mid-Europe. They comprised both
coniferous and broad-leaf species. His data analysis
indicated that when annual N deposition minus an-
nual N increment (N uptake in boles and branches)
was >0, the relationship between this difference and
annual N leaching could be approximated by a 1: 1
line. This conclusion is seemingly a paradox since
microbial or chemical N immobilization is generally
regarded as a key process in the N cycle. Johnson's
hypothesis was that in areas with high N deposition,
soil NH4", i.e. the N substrate for autotrophic nitrifi-
ers, will be continuously kept at a high level. Under
such circumstances the soil nitrifiers seem to be better
competitors for available N compared with trees and
other vegetation. The 1 : 1 relationship implies that
the annual net N mineralization is equal to the N
input in leaf litter plus root litter. Several investiga-
tors have found a close statistical relationship be-
tween N deposition and N leaching, but rather weak
relationships between N deposition and internal re-
cycling processes (Gundersen 1995 and cited refer-
ences). According to Gundersen (1995), changes in
processes such as N mineralization, N immobilization
and N uptake induced by changes in N deposition
may proceed at a much slower rate than correspond-
ing changes in N leaching. In many cases the N
leaching response seems to be more closely related to
hydrological than to biological processes. Cole et al.
(1992) concluded, however, that at some of the sites
discussed by Johnson (1992), N mineralization could
explain more of the variation in NOf leaching than
any of the other single variables evaluated.
In this paper data collected in two Swedish moni-
toring programmes was used to analyse empirically
how N leaching, N immobilization and N uptake by
the trees are related to N deposition. The somewhat
provocative statements made by Johnson (1992) pro-
vided the motivation to write this paper. Special
emphasis is placed on the problem of how the N
immobilization should be treated in large-scale esti-
mates of critical loads of N. A default value (2-5 kg
N ha" 1 yr~') for the- N immobilization was used in
the mass balance calculations from which critical
loads were derived (Downing et al. 1993, Posch et al.
1995). Presently, a number of empirical approaches is
being used in different European countries (Posch et
al. 1997).
Scand. J. For. Res. 13 (1998)
MATERIAL AND METHODS
Data sources
Data were collected from 37 monitoring sites in
forests in south Sweden dominated by Norway spruce
(Picea abies (L.) Karst.) or Scots pine (Pinus sylvestris
L.). Tree mensuration data, i.e. diameter at breast
height (DBH, measured in cm 130 cm above the
ground), tree height and stand age were obtained
from the Swedish National Forestry Board, and data
on bulk deposition, throughfall and ion concentra-
tions in soil water were obtained from a monitoring
programme carried out by one of the authors (O.
Westling) at the Swedish Environmental Research
Institute, Aneboda (Hallgren-Larsson et al. 1995).
Tree growth and N uptake
DBH was measured on all trees in each plot, while
height and age measurements were conducted on a
smaller number of trees. DBH values were used as
input data for the biomass estimates. DBH was con-
verted to biomass (kg dry mass ha~') of individual
tree components, i.e. boles (wood + bark), branches
(wood -I- bark + needles) and stumps plus coarse
roots, by using regression functions for Norway
spruce or Scots pine published by Marklund (1988).
The nitrogen content of the individual tree compo-
nents (kg N ha" 1 ) was then estimated by applying N
concentration values published by Albrektson &
Lundmark (1991). All calculations were based on
DBH measurements conducted in 1984, 1989 and/or
1994. The longest time interval available was used for
the calculations of the annual tree growth during the
last 5- or 10-yr period. Stand age refers to the year of
the first DBH measurements. Tree N uptake was
defined as the summed uptake of all tree components
mentioned above, assuming that they were to be
removed at harvest. Bole N accumulation was the
uptake in bole wood and bole bark, i.e. the biomass
fractions that would be removed at conventional bole
harvest.
Nitrogen deposition and N leaching
Data were collected during at least 2 yrs in the period
1985-1994. Bulk deposition and throughfall deposi-
tion were collected once a month in open funnels
attached to collectors (one collector for bulk deposi-
tion and 10 for throughfall) placed on wooden poles.
During winter, a snowsack and plastic buckets were
used for collecting bulk deposition and throughfall,
respectively. The collected water samples were
Scand. J. For. Res. 13 (1998) Retention of N in coniferous forest ecosystems 395

Table 1 . Summary statistics of the investigated variables

Variable Min. Max. Mean SD

Stand age (yrs) 20 98 62 20

N deposition (kg N ha" 1 yr" 1 ) 4.0 29.4 13.3 • 7.4
N leaching" (kg N h a " 1 yr" 1 ) 0 11.9 0.7 2.1
Tree N uptake * (kg N ha" 1 yr" 1 ) 1.0 27.1 7.6 6.4
Bole N accumulation'' (kg N ha" 1 yr" 1 ) 0.2 13.9 4.0 3.5
Soil N retention1' (kg N ha" 1 yr" 1 ) -2.7 23.2 8.5 6.0
Ecosystem N retention'' (kg N ha" 1 yr" 1 ) 4.0 25.6 12.6 6.3

" Estimated from the chloride mass balance.

h
Summed uptake in boles (bark + wood), branches (bark + wood + needles) and stumps + coarse roots.
' Uptake in bark + wood.
''(Maximum N immobilization), according to eq. (2).
" According to eq. (3).

analysed for pH and major cations and anions.
Amounts of inorganic N, i.e. NH^1" and NOf (kg
N h a " ' yr" 1 ), were obtained by multiplying corre-
sponding water volumes and ion concentrations.
The total N deposition was estimated by adding
bulk deposition and dry deposition. Dry deposition
was obtained according to Lövblad et al. (1992,
1995). The dry deposition estimates were based on
measured concentrations of particulate and gaseous
N compounds for each region and previously estab-
lished deposition velocities in coniferous forests.
Five ceramic suction cups (P80; Staatliche Porzel-
len Manufaktur, Berlin, Germany) were installed at
a soil depth of 0.5 m and used for collecting bulk
samples of soil water on three occasions each year
(March/April, June and October/November). Water
samples were analysed for pH and major cations
and anions. The soil water concentrations used in
this study were median values (for further details,
see Hallgren-Larsson et al., 1995.) Water percola-
tion at 0.5 m soil depth (P; mmH,O) was calcu-
lated by adjusting the measured throughfall
volumes (T; mmH 2 O) for evapotranspiration ac-
cording to
/>=r.Y[Ci-] T /[ci] s w , (i)
where [C1"]T and [Cl~] s w denote the chloride con-
centration in throughfall and soil water, respec-
tively. Chloride was assumed to be a conservative
element which did not interact with the soil matrix.
The resulting water volumes were then used for
converting the N H / and NO3" concentrations in
the soil water to N leaching rates (kg N ha" 1
yr~'). For the sites in south and southwestern Swe-
den, which were the only ones that had measurable
concentrations of mineral N (mostly NO3~), we also
used the S O j " mass balance for comparison, as-
suming a steady state between the SO;j~ input in
throughfall and S O j " leaching. This seems to be a
reasonable assumption for this part of the country
(Hultberg 1985, Hultberg & Grennfelt 1992, Nohr-
stedt et al. 1996).
Two N mass balance equations (eqs 2 and 3)
were used for data interpretation, where N leaching
was estimated from the chloride mass balance:
Soil N retention = N deposition
— bole N accumulation
— N leaching. (2)
Equation (2) shows the estimated soil N retention
(maximum N immobilization) on an annual basis
assuming that only the boles were to be removed
from the site at harvest, which would be the typical
situation in conventionally managed forest stands.
Soil N retention therefore includes N taken up in
branches (including the needles), stumps and roots,
i.e. the N pool which will be incorporated into the
soil organic matter pool for a shorter or longer
period, depending on the degree of remineralization.
The ecosystem retention of N was defined as:
Ecosystem N retention = N deposition
— N leaching. (3)
Budget calculations and statistical calculations were
made using routines in Microsoft Excel (Anon.
1992) and in the SAS statistical package (Anon.
1989), respectively. Average values are presented as
mean values ± SD.
396 S. I. Nilsson et al. Scand. J. For. Res. 13 (1998)

~ 15

10
S?

"5 "10 " 1 5 "20 25 30
N deposition (kg ha"1yr~1)
Fig. I. N leaching versus N deposition. N leaching is
estimated from the chloride mass balance. N leaching =
0.00024 • (N deposition)3, r2 = 0.56, p < 0.0001.
RESULTS
The summary statistics of the investigated variables
are presented in Table 1. Only a few of the investi-
gated stands had a measurable N leaching (Fig. 1).
The mean value calculated for all 37 sites was 0.7 +
2.1 kg N h a " 1 yr~ l (Table 1), while the median value
was 0.05 kg N ha~' yr~'. All stands where N leach-
ing was observed were situated in the southernmost
part of the country. N leaching was positively corre-
lated with N deposition according to the regression
function shown in Fig. 1, although N leaching was
practically zero until the N deposition approached 15
kg N ha~' yr" 1 . There was a reasonably good corre-
spondence between Cl "-based and SOj"-based esti-
mates of water percolation and N leaching (Table 2).
Differences between localities with respect to N leach-
ing were, to a large extent, accounted for by differ-
ences in soil water concentration rather than by
differences in runoff (Table 2).
|
eo
•g ~0 20 40 60 80 100 120
Stand age (yr)
Fig. 2. Bole N accumulation versus stand age. Bole N
accumulation = - 0 . 0 6 • Stand age + 7.71. r2 = 0.10, p<
0.04.
The estimated soil N retention (maximum N im-
mobilization) (eq. 2) was independent of stand age
(p> 0.10), whereas a significant negative correlation
was found for the dependence of bole N accumula-
tion on stand age (Fig. 2). Visual inspection of Fig. 2
indicated that the age dependency was particularly
evident at stand ages 20-40 yrs, which is in accor-
dance with, e.g. Miller & Miller (1988). The soil N
retention according to Eq. (2), increased significantly
with increasing N deposition (Fig. 3). A positive
correlation was also found for the relationship be-
tween bole N accumulation by the trees and N depo-
sition (Fig. 4). Bole N accumulation accounted for
about 53% of the tree N uptake as defined above
(Table 1).
Ecosystem N retention (eq. 3) was positively corre-
lated with N deposition (fl- = 0.92, ^ < 0.0001).
However, N leaching at most of the 37 sites was
approx. zero. Retention values, therefore, became

Table 2. Median NH$ and NOj concentrations, estimates of N leaching based on either the Cl or the
mass balance and water percolation estimates based on the Cl~ mass balance

N leaching

NO3--N NH + -N C l - bal. SO5- bal. C l " bal.

Site (mg N/L) <mg N/L) (kg N ha~ 1 y r - 1 ) (mmH 2 O)

1 0.87 0.01 0.96 1.10 110

2 1.14 • 0.36 3.86' 4.38 258
3 ' 5.79 0.14 11.88 11.88 201
4 0.74 0.01 2.18 2.62 289
5 0.42 0.05 1.59 1.70 344
6 1.25 0.02 2.29 4.09 181
7 2.17 0.03 3.36 5.89 153

Only those sites which had a significant leaching of mineral N were included.
Scand. J. For. Res. 13 (1998) Retention of N in coniferous forest ecosystems 397

Thus, direct estimates of dry deposition, such as

CO
those used in this study, should be preferred. More
JZ
work concerning typical air concentrations and depo-
S sition velocities will have to be done, however, before
c
o these estimates can be used on a routine basis
c
o (Lövblad et al. 1995). Therefore, bulk deposition
I and/or throughfall values are still used in regional
evaluations of the effects of N deposition on other N
CD 0 5 10 15 20 25 30 fluxes in forest ecosystems (Gundersen 1995, Hall-
N deposition (kg ha'1yr~1) gren-Larsson et al. 1995). Alternative deposition cal-
culations were made in which the highest of the
measured NH4" and NOj~ fluxes in bulk deposition
Fig. 3. Soil N retention according to eq. (2) versus N
deposition. Soil N retention = 0.60 • N deposition, r2 = or throughfall deposition at each site were selected
0.63, p < 0.0001. For definitions and explanations, see text (usually the throughfall fluxes). These deposition esti-
and Table 1. mates were approx. 20% lower than those presented
previously which, for instance, caused the estimated
strongly dependent on the actual N deposition values, soil N retention (maximum N immobilization), ac-
implying a non-negligible autocorrelation. Ecosystem cording to eq. (2), to decrease from 8.5 to 5.9 kg N
N retention was negatively correlated with stand age ha" 1 yr~' (cf. Table 1). However, the statistical
(R2 = 0.08, p < 0.05), indicating that most of this age relationships illustrated in Figs 1 - 4 did not change
dependency was accounted for by the bole N accu- with respect to plus or minus signs or with respect to
mulation by the trees (Fig. 2), since the soil N their statistical significance.
retention according to Eq. (2) was independent of
stand age. Soil solution chemistry and water percolation
The N concentrations in the soil solution were only
analysed three times per year, which may have af-
DISCUSSION
fected the accuracy of the estimates. For logistical
Different estimates of N deposition reasons a more intensive sampling scheme was not
The bulk deposition of NOf and, to a lesser extent, feasible. However, the sampling scheme was chosen
NH4" is normally lower than total N deposition, to include contrasting moisture and temperature
particularly in high-deposition areas. The total N regimes (early spring, summer and late autumn),
deposition is not readily obtained from throughfall which were supposed to affect the soil biological
measurements either, because internal N release, or activity and the N concentrations in the soil solution
alternatively, N absorption in the tree canopy will differently. In addition, the median mineral N con-
affect the deposition estimates (Gundersen 1995, centrations used in the N leaching estimates were
Hallgren-Larsson et al. 1995, Lövblad et al. 1995). based on > 2 yrs of data collection. An undue influ-
ence of extreme values (either high or low) is, there-
fore, less likely. For the sites which had a measurable
N leaching according to Table 2, the period of data
collection was > 4 yrs. As for N leaching, differences
between the localities with respect to mineral N con-
c centration in the soil water seemed to be more impor-
.0

tant than differences in water percolation. The good

3 correspondence between calculations based on either
o
Ü
CD
the chloride or the sulphate mass balances indicated
that the water flux estimates were reasonably accurate
5 10 15 20 25 30 compared with other types of estimates (Table 2).
N deposition (kg ha' yr'1)
1 Nohrstedt et al. (1996) found that the water percola-
tion estimates for a coniferous forest stand in south-
west Sweden could vary between 344 mm (model
Fig. 4. Bole N accumulation versus N deposition. Bole N
simulation) and 490 mm (run-off from a nearby
accumulation = 0.21 • N deposition, r 2 = 0.19, p < 0.006.
398 S. I. Nilsson et al.
river). According to chloride and sulphate mass bal-
ances the percolation was 363 mm and 390 mm,
respectively.
N2 fixation, dentrification and leaching of organic N 1937).
A complete input-output budget should include N2
fixation, gaseous losses of N such as denitrification as immobilization) was estimated according to eq. (2) to
well as leaching losses of dissolved organic N (DON). ' be 8.5 ±6.0 kg N ha" 1 yr"1 (Table 1), i.e. 17-43
No information was available from the investigated
sites concerning these processes. They have usually
been regarded as rather insignificant in coniferous
forest ecosystems with acid podsolic soils. Recently,
Sitaula et al. (1995) showed that the N2O flux from a
podzolic soil in Norway, receiving 10-15 kg N ha" 1
yr~ ' as atmospheric deposition, was about 0.4 kg N
ha" 1 yr""1. The N2O flux increased to 0.9 kg ha" 1
yr" 1 when 30 kg N ha" 1 yr"1 was applied as
NH4NO3 fertilizer for 2 consecutive yrs. DON was
not included in the chemical analyses of the soil
water. However, data from two Norway spruce sites,
Hasslöv in south-western Sweden and Strâsan in
south-central Sweden, suggest a concentration in the
order of 0.2-0.5 mg N I"1 as DON in soil water
collected from the lower part, of the B horizon
(Nilsson et al. unpublished data). If annual water
percolation is 200-400 mm, such concentrations
would correspond to an annual leaching of 0.4-2 kg
N ha" 1 . To conclude, ignoring losses of N by deni-
trification processes and leaching of DON probably
resulted in an overestimation of the soil N retention,
1 1
according to eq. (2), of 1-3 kg N ha" yr" .
Long-term ecosystem N retention
The long-term retention of N in Swedish forest soils
since the end of the last glaciation (10 000-12 000 yrs
BP) is in the order of 0.2-0.5 kg N ha" 1 yr"1 (Rosen
et al. 1992). This estimate was based on inventories of
the organic matter content in the mineral horizons of
podsolized forest soils. During the primary vegetation
successions, the N stores of the forest ecosystems
were initially built up by N2-fixing plant species such
as Hippôphaë ramnoides and Alnus spp., in addition
to the N contribution of deposited NH4+ and NOf
(e.g. Tamm 1991). Eventually, the N2-fixing species
decreased in importance owing to increasing interspe-
cific competition and less favourable site conditions,
e.g. decreased soil pH. Nitrogen accumulation was
intermittently interrupted by forest fires. The present
Swedish forests, particularly in southern Sweden,
grow on soils that have been used intensively for crop
cultivation and grazing for hundreds of years. Many
Scand. J. For. Res. 13 (1998)
of these soils should have been depleted of N by
previous management practices, which in many re-
gions included regular burnings to improve the graz-
ing conditions for cattle and sheep (Malmström
The current average soil N retention (maximum N
times higher than the values estimated for the period
since the latest glaciation. Even if denitrification and
leaching of DON arc taken into account, there is
obviously a large discrepancy betweeen the present
estimate and the long-term estimate made by Rosen
et al. (1992). One fundamental difference between our
estimate and that reported by Rosén et al. (1992), is
the time-scale. Much of the organic N that is incor-
porated into the soil organic matter pool on a short-
term annual basis will eventually be remineralized
and taken up by the vegetation. Our estimate seems
more relevant, however, when discussing the ecosys-
tem effects of the current N deposition.
Both maximum N immobilization and N leaching
(Fig. 3, Fig. 1) seemed to be dependent on N deposi-
tion (cf. Gundersen 1995). Nitrogen deposition over
northern and central Europe has increased since the
mid-1950s (Rodhe & Rood 1986, Skeffington &
Wilson 1988). Concerning the time scale, it is interest-
ing to note that N deposition at the Dalby Norway
spruce site in southernmost Sweden (28.8 kg N ha" 1
yr"1) was similar to the N deposition estimated in
1967-1968 (throughfall 4-stemflow amounting to
24.1 kg N ha" 1 yr" 1 ) at a Norway spruce site situ-
ated only about 60 km further north (Nihlgürd 1970).
Eriksson & Johansson (1993) documented that
there has been a significant increase in the volumetric
growth of Norway spruce in south-western Sweden
since the early 1900s. The authors mentioned an
increase in N deposition as one of several possible
explanations for the observed trend. A weak but
significantly positive correlation (p < 0.006) was
found between the estimated bole N accumulation by
the trees and N deposition (Fig. 4), which is partly in
line with the interpretations of Eriksson & Johansson
(1993). The present results differed from those re-
ported by Johnson (1992), who did not find any
relationship between N deposition and N uptake by
the trees, or between N deposition and annual soil N
retention. He demonstrated that soil N seemed to be
extensively depleted in many cases. According to eq.
(2) such a soil depletion was only evident at one of
the 37 investigated sites. One explanation for the
Scand. J. For. Res. 13 (1998)
discrepancies between the two studies could be that
Johnson included the N accumulation in "all woody
tissues" in his calculations, while in the present calcu-
lations, stumps, coarse roots and branches were ex-
cluded. This assumption was tested by calculating: (1)
a linear regression with estimated soil N retention
based on N accumulation in all woody components
versus N deposition, and (2) a separate linear regres-
sion for N accumulation itself versus N deposition. In
both cases positive relationships were still obtained,
although the level of statistical significance decreased
to some extent compared with the analogous linear
regressions based on bole N accumulation, shown in
Figs 2 and 4:
Soil N retention = 0.46 • N deposition; p < 0.0035
N accumulation = 0.36 • N deposition; p < 0.02.
Consequently, a N deposition dependency could still
be demonstrated with respect to both estimated soil N
retention and N accumulation in the trees. Differences
in climate may explain part of the discrepancy between
the two studies. Most of the sites discussed by Johnson
(1992) were situated either in the USA or in mid-Eu-
rope, i.e. at latitudes considerably lower than those
treated in this paper. Average summer temperatures
are higher than in the Swedish stands, implying both
a higher mineralization rate and a higher nitrification
rate. Differences in site history in terms of C and N
net losses might also contribute to the discrepancies
(cf. Aber et al. 1997).
N immobilization and the critical load of N
The mean value of soil N retention according to eq.
(2) (maximum N immobilization) obtained in this
study, 8.5 + 6.0 kg N ha" 1 yr~' (Table 1), was not
very different from the default value range of 2 - 5 kg
N ha" 1 yr" 1 used previously in the international work
on critical loads (Posch et al. 1995) and similar to the
value of 8 kg N h a " ' yr" 1 reported by Staaf et al.
(1997). The present mean value was also close to the
annual soil N retention (5-10 kg N ha" 1 yr" 1 )
estimated for Sitka spruce plantations in Wales receiv-
ing a deposition of 20-30 kg N ha" 1 yr" 1 (Emmett &
Reynolds 1996). Excess N leaching during the clear-
felled stage was partly accounted for by the latter
authors. In many of their investigated stands the
calculated soil N retention based on the N increment
in boles [eq. (2)] was negative before clear-felling (cf.
Johnson 1992, Table 1). Seen over the whole rotation
period, however, there was a net retention.
Retention of N in coniferous forest ecosystems 399
Ecosystem N retention [eq. (3)] showed both posi-
tive and negative values in coniferous forests in conti-
nental Europe, receiving 15-75 kg N ha" 1 yr" 1 as
deposition (Dise & Wright 1995). It is evident that
even in southern Sweden, the leaching situation in
most of the forest stands differs greatly from that in
many other parts of Europe. Dise & Wright (1995)
showed that annual leaching figures in the order of
25-30 kg N were quite common. The highest leaching
in the present study (11.9 kg N ha" 1 yr" 1 ) was
observed at the Dalby Norway spruce site in southern-
most Sweden. However, the soil N retention at the
same site according to eq. (2) was of about equal size
( 12.8 kg N h a " ' yr~ '). Similar leaching rates (10.1 and
12 kg N ha" 1 yr" 1 ) were reported by Bergkvist &
Folkeson (1995) from two nearby sites with Norway
spruce. According to an estimate based on a few case
studies reported by Rosen (1988), the excess N leach-
ing in south Sweden during the clear-felled stage may
not decrease the average soil N retention for the whole
rotation period (70 yrs) by more than 14%. Later
studies of 10 forest sites in southern Sweden showed
a range of 3-34% (Örlander et al. 1997).
Differences in N deposition as well as in climate and
site histories may all contribute to the different N-
leaching conditions in Sweden compared with other
European countries. Nevertheless, there is every rea-
son to believe that N saturation will eventually be
reached in some of the forest stands in southern
Sweden. Before accurate predictions can be made
concerning the future N status at either present, higher
or lower N deposition levels, more detailed experimen-
tal work will have to be conducted on some of the key
processes of the N cycle, such as N mineralization, N
immobilization and nitrification. Presently, qualitative
indices, such as needle N content the needle arginine
content, or the C/N ratio in different soil layers, can
only be used to predict the likelihood of nitrification
and NO3~~ leaching (cf. Nohrstedt et al. 1996). In
modelling work, process descriptions based on sound
and widely accepted plant physiogical and soil biolog-
ical concepts have to be used. The previously men-
tioned Soil N and PnET-CN models look promising
in this respect, provided that there is continuous
feedback between experimental work and modelling
work.
ACKNOWLEDGEMENTS
This study was funded by the Swedish Environmental
Protection Agency. Tree diameters and tree heights
400 5. /. Nilsson et al.
were provided by the Swedish National Forestry
Board (through Sören Berghäll). Anna Hedlund car-
ried out most of the computer work and Mats Linde
provided constructive criticism. David Tilles signifi-
cantly improved the English.
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