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Impact of dolomite lime on the ground vegetation and on potential net N transformations in Norway spruce
(Picea abies (L.) Karst.) and sessile oak (Quercus petraea (Matt.) Lieb.) stands in the Belgian Ardenne
Jean-François Dulière, Monique Carnol, Shanti Dalem, Jean Remacle, François Malaisse
To cite this version:
Jean-François Dulière, Monique Carnol, Shanti Dalem, Jean Remacle, François Malaisse. Impact of
dolomite lime on the ground vegetation and on potential net N transformations in Norway spruce
(Picea abies (L.) Karst.) and sessile oak (Quercus petraea (Matt.) Lieb.) stands in the Belgian
Ardenne. Annals of Forest Science, Springer Nature (since 2011)/EDP Science (until 2010), 1999, 56
(5), pp.361-370. �hal-00883279�
Original article
Impact of dolomite lime on the ground vegetation and
on potential net N transformations in Norway spruce
(Picea abies (L.) Karst.) and sessile oak (Quercus
petraea (Matt.) Lieb.) stands in the Belgian Ardenne
Jean-François Dulière Monique Carnol Shanti Dalem
Jean Remacle François Malaisse
a
Faculté des sciences,
biologie végétale,
université de Mons-Hainaut, avenue Maistriau, 23, 7000 Mons,Belgium
"Laboratoire
d’écologie,
faculté universitaire des sciencesagronomiques,
passage desdéportés,
2, 5030 Gembloux,Belgium
c
Département
debotanique
B22,écologie
microbienne etradioécologie,
université deLiège,
Sart Tilman, 4000Liège, Belgium
(Received 15
September
1998;accepted
25 November 1998)Abstract - The
impact
of dolomite lime (5T·ha -1 )
on theground vegetation
and onpotential
netnitrogen
(N) transformations wasinvestigated
in twoBelgian
forest ecosystems.Norway
spruce (Picea abies (L.) Karst.) and sessile oak (Quercus petraea (Matt.) Lieb.) stands were situated in the Haute Ardenne (eastBelgium)
on acid-brown soil. Theherb-layer
floristic richness increased dur-ing
the 2 yearsfollowing liming,
with the appearance oflight
andN-demanding species,
which are also found in clear-cut areas or onroad verges. Mosses reacted
rapidly, showing
a decreaseacidophilous-dominant species
and the establishment of some ruderalspecies.
Six months afterliming,
thepH
wassignificantly
increased in theorganic
horizon of both stands and in theorganomineral
horizon of the oak stand. Soils
originating
from the two stands showed distinct responses in netNO 3 - production
to the dolomite lime treatment. In theorganic layer
of the Quercus soil, netNH 4 + production
was decreased,NO 3 - production
increased, and total N min- eralisation remainedunchanged.
In theorganomineral layer, NO 3 - production
was increased. In the Picea soil,NO 3 - production
wasdecreased in the
organomineral
soillayer.
These results indicate thepossibility
of differences in the control of the N transformation processesoccurring
in the two sites. (© Inra/Elsevier, Paris.)dolomite
liming
/ forest / N mineralisation / nitrification /ground vegetation
Résumé - Effet d’un amendement
calcaro-magnésien
sur lavégétation
et les transformationspotentielles
nettes de l’azotedans une
pessière
(Picea abies (L.) Karst.) et une chênaie(Quercus
petraea (Matt.) Lieb.) en Ardennebelge. L’impact
del’apport
de 5 T ha-1 de dolomie sur lavégétation
et lecycle
de l’azote a été étudié dans deuxécosystèmes
forestiers, situés en Ardennebelge,
uneplantation d’épicéas
(Picea abies (L.) Karst.) et une chênaie à Quercus petraea (Matt.) Lieb. La strate herbacée s’enrichit lors des deux annéesqui
suivent l’amendementd’espèces pionnières
ounitrophiles caractéristiques
des trouées ou desbords de chemins. La strate muscinale
réagit rapidement
par larégression
desespèces acidophiles
dominantes etl’apparition
discrètede
neutrophiles
ou de rudérales. Six moisaprès
l’amendement, lepH
aaugmenté significativement
dans l’horizonorganique
desdeux
plantations
et dans l’horizonorgano-minéral
de la chênaie, Laproduction
nette deNO 3 -
du sol a été influencée différemment par l’amendement dans les deux sites. Dans l’horizonorganique
de la chênaie, laproduction
nette deNH 4 +
a été diminuée, laproduc-
tion de
NO 3 - augmentée,
sans modification de la minéralisation totale. Dans l’horizonorgano-minéral,
laproduction
deNO 3 -
a aug-menté dans la chênaie, alors
qu’elle
diminuait dans lapessière.
Ces résultatsindiquent
lapossibilité
de contrôles différents des trans- formations d’azote dans les deux sites. (© Inra/Elsevier, Paris.)amendement
calcaro-magnésien
/ forêt / minéralisation / nitrification /végétation
*
Correspondence
andreprints
jean-francois.duliere@umh.ac.be
1. Introduction
Forest decline
symptoms
observed inEurope
since theearly
1980s have affectedBelgian
forestsparticularly
forNorway
spruce(Picea
abies(L.) Karst.)
and oaks(Quercus
petraea(Matt.)
Lieb. andQ.
roburL.).
Insouthern
Belgium,
theproblem
isbeing
studiedby
sever-al
university
teams, in aninterdisciplinary
programmefinanced
by
the Walloonregion (Section
Nature andForestry) [17].
Many
biotic and abiotic factors govern forestdieback,
the extent of which oftendepends
onregions
and treespecies.
Soil acidification due toatmospheric pollution,
and
subsequent
nutritionaldeficiencies, appeared
to be amajor
cause of the observed decrease in forest health.This was
especially
confirmed forNorway
spruce grow-ing
onnaturally
poor acidic soils of theBelgian
Ardenne[35, 36].
ForQ.
robur andQ.
petraea, many studies revealed that the decline should be considered as a com-plex-causal phenomenon [15, 16, 23]. Among predispos- ing factors,
nutritional deficiencies can weaken the trees, which are less able to support further stress. Fertilisation is thensuggested,
with a view toreplenish
the low level of some elements and to restore a nutritional balance consistent with therequirement
of treespecies [18].
Magnesium (Mg) deficiency
is oftenpointed
out as amajor
cause of decline in hardwood and coniferous forests[7, 20, 28, 34].
With the aim to raise lowpH
andto
supply
deficientMg,
dolomite lime is oftensuggested.
In
addition,
the calcium andMg supply
may reduce alu- minium in the soil cationexchange complex [26].
However, increasing
soilpH
could lead to increased nitrate and associated cationleaching [21 ].
In western
Europe,
many scientific research teamshave
investigated
the effects ofliming
on different parts of forestecosystems.
Theimpact
on health conditions andgrowth
oftimber-producing species
hasfrequently
been studied
[5, 6, 25, 26].
The reaction ofground
florato
liming
has sometimes received attention[19, 27, 29, 31, 33]. However,
theimpact
on mosses hasrarely
beeninvestigated quantitatively [1].
We
present
astudy
on the modifications inducedby
adolomite lime treatment on
neighbouring Norway
spruce and sessile oak forest stands. We focus on the botanicalaspects,
soil chemical parameters andpotential nitrogen (N)
mineralisation rates.2. Materials and methods
2.1. Location and
experimental design
The site was situated in the
’Hertogenwald’
Forest(50°34’ N,
6°02’E),
easternBelgium,
at 440 m in alti-tude and with an annual rainfall of about 1 150 mm and a mean annual air temperature of 8.1 °C
(source:
Meteorological Royal
Institute ofBelgium).
The soil is
acid-brown,
derived from aprimary
Revinien
quartzitic substrate,
with whiteclay occurring
at about 30 cm in
depth.
Twoneighbouring
stands with moder todysmoder
humus type and characterisedby
thepresence of
pseudogley
were studied. The P. abies stand is secondgeneration, planted
in 1930. Theoriginal
vege- tation was a mosaic of deciduous forest and moorland.The
ground vegetation
isscattered,
except in gaps where Moliniacaerulea,
Pteridiumaquilinum
andDeschampsia flexuosa essentially
are more abundant.The moss
layer
is welldeveloped,
with various Dicranaceaespecies
andPolytrichum formosum
as dom-inant taxa. The
Q.
petraea standoriginates
from a cop-pice (with Fagus
sp. and Betulaspp.) dating
fromapproximately 1930,
when oaks were favoured. The herblayer
is dominatedby
M. caerulea and P.aquilinum,
with
essentially
D.flexuosa,
Vacciniummyrtillus
andCarex
pilulifera.
Thisvegetation
can beregarded
as aLuzulo-Quercetum molinietosum, according
to Noirfalise[24]. Throughfall
Ninputs (under Picea)
are about 20 and 15kg·ha -1 ·year -1 NH 4 + -N
andNO 3 - -N, respectively.
Twelve square
plots
of 225m 2
were established in each forest type around a central dominant or co-dominant tree, selectedrandomly.
A minimum of 5 m between eachplot
was
respected
toprevent
cross-contamination. Sixplots
ofeach stand were limed in
April
1996(figure 1)
with 5T·ha
-1
of a dolomite limesuspension (55/40).
To ensurehomogeneity
of lime distribution within theplots,
it wasapplied manually
with aportable spraying equipment,
dis-pensing
thesuspension
at a constant rate[ 14].
2.2. Soil chemical characteristics
Soil
samples
were taken from eachplot
6 months afterliming.
Theorganic (4-8
cm inheight) layer
and the first 10 cm of theorganomineral layer
wereseparated
foranaly-
ses.
pH
was measuredpotentiometrically
on fresh soil in1:2
(v/v) suspensions
in demineralised water.P, K,
Ca andMg
were measured at the ’Stationprovinciale d’analyses agricoles’
of Tinlot(Belgium). Exchangeable
cations weremeasured
by
the CSW-EDTApH
4.65 method and spec- trometric atomicabsorption. Phosphorus
was extractedwith citric acid and measured
by colorimetry.
2.3. Potential N mineralisation
Two intact soil cores
(PVC,
9.5 cmdiameter)
contain-ing
a 15-cmlength
of soil were taken from eachplot
inOctober
1996,
at 1-3 m around the central tree.Samples
of one
plot
were taken close to eachother,
to minimisespatial variability.
One core perplot
wasimmediately analysed
for mineral N content andpH (see earlier).
Thesecond core was incubated for 60 d in the
laboratory
inthe
dark,
at 80 % fieldcapacity (100
% fieldcapacity
was defined as the water
remaining
after a water-saturat-ed core was allowed to drain for 12
h)
and 20 °C. Thewater content was
adjusted
every 4 d with distilledwater.
For
analyses,
the cores were divided intoorganic (comprising Ol,
Of andOh)
andorganomineral (Ah;
subsequently
calledmineral) layers,
andweighed. They
were
homogenised manually.
The water content wasdetermined as
weight
loss at 105 °C.Exchangeable NH 4 + -N
andNO 3 - -N
wereanalysed
after extraction
(1 h)
with 125 mL KCl 6 % of 20 gorganic
soil and 50 g mineral soil[2],
followedby
steamdistillation of 20 mL of filtered extract. In a first
step,
ammonia was liberated from the extract in the presence ofMgO
and collected in a vesselcontaining
boric acidcombined with an indicator solution. In the
remaining
filtered extract,
NO 3 - -N
was reduced toNH 4 + -N
in thepresence of Dewarda’s
alloy,
distilled and collected in asecond
receiving
container.NH 4 + -N
was thenanalysed
by
titration with 0.005 NH 2 SO 4 [8].
Previousanalyses
had shown
NO 2 - -N
concentrations to beinsignificant [9].
Net N
mineralisation,
ammonium and nitrateproduc-
tion are
expressed
as the difference between contents after and before incubation.They
areexpressed
as mg Nper 100 g
dry weight produced during
60 d. The use ofcores of a known diameter allowed
productions
on anareal basis to be calculated and data for 60 d were multi-
plied by
6.08 toprovide
annual estimates. Within thetwo stands
significant
differences between limed and controlplots
wereanalysed
with a t-test[30].
2.4. Ground
vegetation
surveyThe herb
layer
was listedusing
theBraun-Blanquet
method on the 13 x 13 m inside surface of each
plot (leaving
a 1-m borderalong
theplot boundaries).
Quantitative
data wereprovided using
a’point-intercept’
method
[22]
and will be described later.The moss
layer
survey was conducted on 25 small permanentquadrats
of 50 x 50 cm,systematically
installed on the soil surface in each
plot. Frequency
ofmoss and liverwort
species
in aplot
was estimatedby
thenumber of
quadrats (from
a total of25)
where thespecies
occurred. Afrequency
index was then affected to thespecies,
as follows: 1 =species present
in 1-5quadrates
in theplot;
2 =6-10;
3 =11-15;
4 =16-20;
5 = 21-25.
Data
concerning bryophytes
onstumps
and trunkswere also collected.
3. Results and
discussion
3.1. Soil chemical characteristicsThe dolomite lime treatment resulted in a
significant (P
<0.05) pH
increase in bothlayers
of theQuercus
soil(table I). pH
in water was increasedby nearly
1 unit inthe
organic layer (pH H2O 5.2).
In the minerallayer,
thepH
increase was lesspronounced
but stillsignificant.
The
pH
increase in theorganic layer
of the Picea standwas similar to that of the
Quercus
stand. In the minerallayer
of thisstand, pH
did notchange significantly.
Thetreatment
rapidly
increased theexchangeable
Ca andMg
contents of the
organic
andorganomineral layers
in bothQuercus
and Picea stands. The other essential elements remainedunchanged,
except for P which increased in theorganic layer
of the limedplots
underQuercus.
These results also
clearly
show that the Ca content of the minerallayer
in controlplots
of bothQuercus
andPicea soils should be considered as deficient or at least
not
optimum, according
to the limit of 30mg·100 g -1
suggested by
various authors(e.g. [11]).
3.2. Potential N mineralisation
Potential net N transformations were affected differ-
ently by
the lime treatment in the two stands(figure 2).
Net
NO 3 - -N production significantly
increased in theorganic layer
of theQuercus soil,
with asignificant
reduction in
NH 4 + -N production.
Total net N mineralisa- tion was not affected. In the minerallayer
of theQuercus soil,
the netNO 3 - -N production
alsosignificantly
increased. In the Picea
soil,
a decrease in the netNO 3 - -N
production
in the mineral soillayer (P
=0.07)
was theonly significant
effect. These resultsclearly
demonstrate that moderate doses of dolomite lime(5 T·ha -1 )
canmodify potential
net N transformations in the forest soil of theBelgian
Ardenne.Responses
differed between twoadjacent plots
with similar humus form and on the sameparent material. Six months after
liming, potential
netNO
3 -
-N production
increased in theorganomineral lay-
ers of soil
originating
from aQ.
petraeastand,
whilst adecrease was observed in the mineral soil of a P. abies soil. Increased nitrification without an increase in miner- alisation has been
reported
foroak, Douglas
fir and Scotspine
stands in the Netherlands[13].
In contrast, increased net N mineralisation without an increase in theproportion
of N nitrified wasreported
for asandy
Scotspine
soil[3].
Kreutzer[21]
also found increased nitrifica-tion,
but located in the minerallayer
and linked toincreased mineralisation.
However,
it has to bekept
inmind that these results
apply
6 months afterliming,
andcould
change
in thelong
term. Inparticular,
future sam-plings
will determine whether adelayed
response in thepotential
nitrification to theliming
treatment occurs inthe P. abies stand.
The increase in nitrification indicated the presence of acid-sensitive
chemolithotrophic
bacteria in bothlayers
of the
Quercus
soil. We cannot exclude thepossibility
ofacid-sensitive nitrification in the Picea
soil,
because thepH
increase due toliming
wasrelatively low,
and untilnow restricted to the
top
3 cm of theorganic layer (L.
Ruess, personal communication). However,
the decrease in netNO 3 - -N production
in the minerallayer
indicatedthat different N
cycling
processes andstrategies might
operate in the Picea soil. A watershed
liming experiment (Picea,
3T·ha -1 dolomite)
also showed no effect on soil solution and stream waterNO 3 concentrations, despite
apH
increase in the upperlayer
soil solution[10].
Sufficient N
supply,
the presence of a morhumus,
a C/N ratio(Oh)
below 28 and aeration have been cited asfactors
favouring
increasedNO 3 - -N
lossesfollowing liming [21].
Belkacem andNys [4] compared
responsesto lime of mull
(oak)
and moder humus(spruce) types,
and
reported
arelatively higher
increase in nitrification in the mull humus. Soils used in ourstudy
both had amoder
humus,
but theorganic layer
was thinner in theQuercus
soil. This could indicate better nutrientcycling conditions,
as alsosuggested by higher
nutrient concen-trations and
pH
beforeliming. Similarly,
treedensity
inthe field is more than double in the Picea stand
[14], possibly leading
to differenttemperature
and moisture conditions with different bacterialpopulations.
Furthermore,
it should be noted that we measured netfluxes, possibly resulting
fromchanges
in microbial N assimilation[32].
Potential N mineralisation rates were of the same
order of
magnitude
in bothsoils,
but under control con-ditions,
net nitrification washigher
in the Picea stand.This and the different responses to lime could indicate nitrification to be acid-sensitive in the
Quercus
stand andacidophilic
in the mineral soil of the Picea stand. Even if the bulk soilpH
wasunchanged
in thislayer,
micrositeconditions
might already
have been modifiedby
the lim-ing operation.
Differences in acidsensitivity
of nitrifiers in litter or humuslayers
have beenreported by
De Boeret al.
[12].
Furtheranalysis
of theorganisms responsible
for nitrification in these soils and their
ecological requirements
would lead toimproving
ourunderstanding
of the nitrification process in acid forest soils.
Under control
conditions,
annualpotential
net nitrifi-cation was
highest
in theorganic layer
of the Piceasoil,
where it reached 126
kg·ha -1 ·year -1 NO 3 - -N; however, variability
amongplots from
the same stand washigh (table II).
In theorganic layer
of theQuercus soil,
pro- duction wasonly
16kg·ha -1 ·year -1 NO 3 - -N.
In bothstands,
netNO 3 - -N production
was lower in the minerallayer. Liming significantly
increased netNO 3 - -N
pro-duction in the
organomineral layers
from theQuercus plot.
ReducedNH 4 + -N production (P
<0.1)
was observed in theorganic layer
ofQuercus soil,
and increasedNH 4 + -N production
in the minerallayer
of thePicea soil.
3.3. Ground
vegetation
A
rapid
reaction of the herblayer
toliming
wasobserved, essentially
underNorway
spruce. As shown in tables III andIV,
1 year after treatment thespecies
diver-sity
increased in limedplots, owing
to the emergence ofseedlings belonging
topioneer species (Salix
caprea, Seneciosylvaticus),
orlight
andN-demanding
ruderals(Epilobium angustifolium,
Taraxacum sp.,Epilobium
montanum, Urtica
dioica,
Cerastiumfontanum subsp.
vulgare,
Stellariamedia).
Several(S.
capraea, Taraxacum sp., E.angustifolium)
hadalready appeared
in spruce
plots
ingreat
numbers in summer1996, only
afew months after treatment
(figures
3 and4),
demon-strating
a greatcapacity
torespond
to soil disturbances.This colonisation of limed
plots by nitrophilic species
has also been observed some years after treatment
[27]
as well as in a
long-term
survey[19].
The reaction in oakplots
was lessspectacular, owing
to agreater competi-
tion for new
seedlings by
M. caerulea and P.aquilinum,
and a
quite
thicklayer
ofnon-decayed
and stratified lit- ter, unfavourable to the emergence of young shoots.In both
stands,
the initial vascularvegetation
did notseem to be affected
during
the 2 yearsfollowing
thetreatment. The dominant
species,
M.caerulea,
P.aquil-
inum, D.
flexuosa,
V.myrtillus
and C.pilulifera,
did notshow any
sign
of extension orregression.
Theapparent
difference for somespecies,
such as D.flexuosa,
between limed and control
plots (see
tableIV),
was notdue to treatment, but to an
original heterogeneity
between
plots,
asproved by
pretreatment observations(not presented here).
Aprevious liming experiment
withCa and
Mg
also showed no influence on the behaviour of Molinia and Pteridium[33].
Furtherquantitative
obser-vations should
give
us more informationduring
the com-ing
years.Concerning
the mosslayer,
the dominantspecies
under the spruce cover,
essentially Dicranaceae,
werelargely
affectedby
the treatment(table V).
Thefrequen-
cy of
Campylopus flexuosus,
Dicranum montanum,Dicranella
heteromalla,
forexample,
wasclearly
reduced. At the same
time,
we noted the emergence or the extension of feweracidophilous species (Brachythecium rutabulum, Eurhynchium praelongum)
or
ruderals,
to a minordegree (Ceratodon
purpureus, Funariahygrometrica, Bryum
argenteum,Bryum
rubens).
It isinteresting
to note that thesespecies
alsocover the areas that have been used to clean the material after the
liming operation,
and therefore areimproved
indolomite. The behaviour of the other
species
could notbe
clearly
deduced from these firstinvestigations,
whichstill agrees with a mid-term survey in similar conditions
[1].
We can therefore expect that the immediate reaction of mosses will still be morepronounced
in thecoming
years.
Moreover, although
theglobal
number ofbryophyte species
was notsignificantly altered,
theirground
coveragedecreased,
with the decline of the mostabundant
species.
This isparticularly
evident in the Piceastand,
where the mosslayer
wasimportant.
4.
Conclusion
First
results,
6-18 months after treatment, demonstrat- ed a difference in theimpact
of dolomite lime onadja-
cent spruce and oak
plots
on acid-brown soils withquite
similar chemical soil characteristics. Whereas
pH
andpotential
nitrification weremostly
affected in the oakstand,
herbs and mosses wereparticularly
influenced in the spruce stand. Potential net nitrateproduction
andpH
were increased up to 15 cm in
depth
in the oakplots.
Inthe spruce
plots, pH only
increased in the upperlayer
and net nitrate
production
decreased(P
<0.1)
in theorganomineral
horizon. The appearance ofN-demanding
herbs and less
acidophilous
mosses in the sprucestand, however,
indicated thatchanges
in thebiogeochemical cycling might
have been causedby
the dolomite limetreatment. Results so far demonstrated the immediate
impact
offorestry
managementpractices
notonly
on soilchemistry
but also on thenon-woody
forestecosystem components.
Future data will show the relevance of ourresults for
long-term effects,
inparticular
whether the differentimpact
of lime in the two ecosystems will be exacerbated ordisappear.
Acknowledgements:
Thisstudy
was conducted with thesupport
of the Fonds national de la recherche scien-tifique
ofBelgium.
Thanks to Dr H.Stieperaere
for con-firmation of some
bryophyte identifications,
and to A.Piret for
logistic help.
We would like to thank the Division Nature et Forêts of the Walloonregion
forfinancial assistance for
fencing
theplots,
and forestersand staff for assistance
during
the installation of theexperiment.
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