Biologia 68/6: 1193—1197, 2013
Sec ion Zoology
DOI: 10.2478/s11756-013-0261-8
Respi a ion o wood an nes ma e ial affec ed by ma e ial
and o es s and cha ac e is ics
Ve onika Jilko a1,2,TimoDomisch3,4,ZuzanaHo icka1,2 &JanF ouz1,2
1Ins i u e o Soil Biology, AS CR, Na Sadkach 7,CZ-37005 Ceske Budejo ice, Czech Republic;
e-mail: jilko a. e [email protected]
2Ins i u e o En i onmen al S udies, Cha les Uni e si y in P ague, Bena ska 2,CZ-12800 P aha, Czech Republic
3Facul y o Fo es y, Uni e si y o Joensuu, P.O. Box 111,FI-80101 Joensuu, Finland
4Finnish Fo es Resea ch Ins i u e, Joensuu Resea ch Uni , P.O. Box 68,FI-80101 Joensuu, Finland
Abs ac : We s udied diffe ences in espi a ion o ma e ials om diffe en pa s o wood an nes ( op, bo om, and im)
and om he nes su oundings (humus laye and mine al soil). Samples we e aken om 8 wood an (Fo mica aquilonia)
nes s in each o he wo ypes o o es (bi ch and pine) in eas e n Finland. The diffe ences we e ela ed o ma e ial and
o es s and cha ac e is ics (i.e., mois u e, pH, ca bon con en , and C:N a io). As a esul , he highes espi a ion pe g
DW was measu ed a he op o an nes s in he bi ch o es . Howe e , espi a ion did no significan ly diffe be ween he
pa s o an nes s in he pine o es . Respi a ion o he humus laye s in bo h o es s ands was on a e age highe , whe eas
espi a ion o he mine al soils in bo h o es s ands was lowe in compa ison wi h espi a ion o he nes ma e ials. The
espi a ion pe g C did no show any significan diffe ences be ween diffe en pa s o nes s and su ounding soil. The mos
impo an ac o s influencing espi a ion o he ma e ials appea ed o be mois u e, ca bon con en , and pH. In conclusion,
espi a ion o wood an nes ma e ial is affec ed by he specific ma e ial and o es s and cha ac e is ics.
Key wo ds: Fo mica aquilonia; bi ch o es ; pine o es ; CO2; mois u e; ca bon con en ; pH
In oduc ion
Wood an s (Fo mica s. s ., Hymenop e a: Fo micidae)
include species o an s ha a e dominan in empe a e
and bo eal o es ecosys ems (Dlusskij 1967). They a e
widesp ead om he empe a e zone o Eu ope and Asia
o he No h-Palaea c ic a ea, e en no h o he A c ic
Ci cle (Czechowski e al. 2002; Pun ila & Kilpel¨ainen
2009). Wood an species a e mainly associa ed wi h
coni e ous and mixed o es s, al hough hey occu also
in deciduous o es s (Czechowski e al. 2002).
Due o hei o aging and building ac i i ies hey
affec especially nu ien cycling (F ouz e al. 2005;
F ouz & Jílko á 2008). Recen ly, hei nes s we e ound
o be “ho spo s” o CO2p oduc ion (Ohashi e al.
2005; Risch e al. 2005; Domisch e al. 2006). This
CO2o igina es om se e al sou ces: an espi a ion,
decompose communi y espi a ion, and espi a ion o
oo s pene a ing in o he belowg ound pa s o he nes
(Lenoi e al. 2001; Risch e al. 2005). In ou s udy, we
ocused on he decomposi ion ac i i y o o ganisms (i.e.,
CO2efflux), which is an impo an pa o he ca bon
cycle (S eubing 1970). The knowledge o CO2sou ces
and hei dependence on en i onmen al cha ac e is ics
is impo an o he unde s anding o ecosys em ca bon
balance in o es ecosys ems.
Respi a ion o he decompose communi y can be
influenced by many ma e ial p ope ies, such as mois-
u e, acidi y, ca bon con en , C:N a io, o he ela-
ion be ween cellulose and lignin (Edwa ds e al. 1970;
S eubing 1970; Paul & Cla k 1996). Al hough he an
nes ma e ial is epo ed o be d ie in compa ison
o he nes su oundings, decomposi ion p ocesses a e
a he a ou ed he e (F ouz e al. 1997; Laakso &
Se ¨al¨a 1997; Lafleu e al. 2002). Since he wa e con-
en diffe s be ween he pa s o he nes , he mois u e
is highes in he uppe pa o he nes whe e he e was
also measu ed he highes empe a u e (Coenen-S aß e
al. 1980). Thus, mic obial ac i i y and decomposi ion
p ocesses a e encou aged especially in mois su ace lay-
e s o an nes s (Laakso & Se ¨al¨a 1998; Daube e al.
2001; F ouz e al. 2003). Wood an nes s and hei close
icini y a e usually epo ed o be less acidic in compa -
ison wi h he wide su oundings (Jílko á e al. 2011,
2012), which could also a ou he ac i i y o bac e ia
and ac inomyce es, no only ungi ypical o acidic soils
(Paul & Cla k 1996). Because o o aging and building
ac i i ies o an s, amoun o o ganic ma e (i.e. ca bon
con en ) is enhanced in an nes s (F ouz e al. 1997),
bu he quali y o o ganic ma e (C:N a io and he
ela ion be ween cellulose and lignin) depends on he
sou ce o o ganic ma e and hus on he o es s and
(Edwa ds e al. 1970).
In his s udy we examined ma e ial espi a ion in
diffe en pa s o an nes s in wo o es s ands and in
he humus laye and mine al soil om he su oundings
c
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1194 V. Jilko a e al.
Table 1. P ope ies o ma e ials om he espec i e sampling loca ions a he bi ch and he pine o es .
Sampling Mois u e (%) pH Ca bon con en (%) C:N a io
loca ion F(4,56) =5.04P<0.05 F(4,56) =6.16P<0.001 F(4,56) =6.79P<0.001 F(4,56) = 15.76 P<0.001
Bi ch Pine Bi ch Pine Bi ch Pine Bi ch Pine
TOP 41.15 ±1.85a 19.55 ±1.40w 4.92 ±0.14a 4.50 ±0.05w 43.08 ±0.71a 52.30 ±1.11w 20.97 ±0.13a 67.70 ±6.08w
BOTTOM 22.50 ±1.43b 16.80 ±1.33w 5.23 ±0.08a 5.19 ±0.05x 32.58 ±0.88b 25.40 ±5.44x 19.22 ±0.78a 30.27 ±2.81x
RIM 21.80 ±5.00b 11.46 ±1.22w 5.24 ±0.07a 4.86 ±0.06y 12.07 ±4.14c 9.14 ±0.26y 19.05 ±0.42a 30.12 ±0.15x
HUMUS 59.01 ±2.18c 51.19 ±4.97x 3.97 ±0.07b 3.73 ±0.07z 45.44 ±1.01a 29.74 ±2.55x 22.50 ±0.80a 31.23 ±0.35x
SOIL 16.15 ±1.82b 20.00 ±0.75w 4.81 ±0.12a 5.18 ±0.03x 0.63 ±0.03d 2.23 ±0.16y 30.78 ±3.08b 36.07 ±1.46x
Explana ions: TOP e e s o he op o he an nes s, BOTTOM o he base o he nes s, RIM o he pe iphe y o he nes s, and
HUMUS and SOIL o he humus laye and mine al soil, espec i ely, as con ols. Mean ±SEM and esul s o h ee-way ANOVAs a e
shown. S a is ically homogeneous g oups o alues o he same ma e ial p ope y a e ma ked by he same le e (one-way ANOVA,
Tukey HSD pos hoc es ).
as con ols. We ela ed he diffe ences o mic osi e and
o es s and cha ac e is ics. Ou hypo heses we e ha
(1) ma e ial espi a ion is he highes in he op o he
nes s, (2) he diffe ences in ma e ial espi a ion a e ex-
plained by ma e ial p ope ies (e.g., mois u e, pH, ca -
bon con en ), and (3) he o es s and cha ac e is ics
a e esponsible o he diffe ences in ma e ial espi a-
ion, especially due o ca bon con en o he li e .
Ma e ial and me hods
S udy si es
The s udy was conduc ed in Augus 2009 in wo o -
es s ands, bi ch (Be ula pendula Ro h) and pine (Pinus
syl es is L.), in eas e n Finland nea Joensuu (62◦3928 N,
29◦3938 E, bi ch s and; 62◦4120 N, 29◦4433 E, pine
s and). The bi ch o es was a 47-yea old s and wi h a ee
densi y o 2,376 ees pe ha and an an nes densi y o 9.9
nes s pe ha. An nes s in he bi ch o es we e cons uc ed
mainly om bi ch wigs, ba k, and lea es. Howe e , a small
po ion o he pine needle ma e ial was also included since
he bi ch o es s and was si ua ed ca. 100 m om he pine
o es s and. The pine o es was a 37-yea old s and wi h
a ee densi y o 2,235 ees pe ha and an an nes den-
si y o 6.3 nes s pe ha. An nes s in he pine o es we e
cons uc ed mainly om pine wigs and needles, wi h a con-
side able amoun o coni e esin.
Sampling design
Samples o ma e ials we e aken wi h a sho el om 8
mounds o Fo mica aquilonia Ya ow, 1955 an s in bo h
o es s ands. Sampling loca ions we e he op (20 cm be-
low he su ace), he bo om ( he cen e o he mounds on
he le el o he o es floo ) and he soil im o he mounds
( he pe iphe y o he mounds), and he humus laye and
mine al soil app oxima ely 10 m om an adjacen mound
as con ols. In o al, 80 ma e ial samples we e aken. F esh
samples we e immedia ely anspo ed o he labo a o y o
analyses.
Ma e ial analyses
Respi a ion o he ma e ials, mois u e, pH, and con en o
ca bon and ni ogen we e measu ed. Respi a ion was es-
ablishedusing5go he eshma e ialsa e a wo-day
incuba ion a 20◦
C by i a ion o NaOH acco ding o Page
(1982). Mois u e was de e mined as a diffe ence be ween
esh mass and d y mass a e d ying a 105◦
C o 12 h.
pH was measu ed in a 1:10 ma e ial:wa e suspension wi h
a glass elec ode. The o ganic ma e con en was assessed
based on loss-on-igni ion a e 5 h in 600◦
C. The con en o
ca bon and ni ogen was analyzed in samples o d y c ushed
soil using elemen al analyze EA 1108 (Ca lo E ba Ins u-
men s).
S a is ical analyses
Th ee-way ANOVA wi h o es s and, sampling loca ion and
nes as ca ego ical a iables, wi h nes as a andom effec
nes ed in o es s and, we e used o analyze diffe ences in
ma e ial p ope ies. I he in e ac ion be ween o es s and
and sampling loca ion was significan , one-way ANOVA and
pos hoc es (Tukey HSD) was pe o med o assess di -
e ences be ween he sampling loca ions. All compu a ions
we e made using he p og am S a is ica 10.
Resul s
Ma e ial espi a ion pe g DW soil diffe ed signifi-
can ly be ween he sampling loca ions and o es s ands
(Fig. 1). Diffe ences we e mo e p onounced in he bi ch
o es han in he pine o es . The highes espi a ion
was measu ed a he op o an nes s in he bi ch o -
es , whe e i was se en imes highe han espi a ion
a he op o an nes s in he pine o es . The bo om
and im in bo h o es s ands, as well as he op o
an nes s in he pine o es we e no ma kedly diffe en
in ma e ial espi a ion. Respi a ion o he humus lay-
e s in bo h o es s ands was significan ly highe han
espi a ion o mine al soils; i was also highe han es-
pi a ion a he bo om and im o an nes s in bo h
o es s ands, whe eas espi a ion o he mine al soils
was lowe . When we calcula ed espi a ion pe g C, he
diffe ences be ween indi idual pa s o nes s and nes
su oundings we e no significan .
The o he ma e ial p ope ies also conside ably di -
e ed be ween he wo o es s ands (Table 1). An nes s
in he bi ch o es we e on a e age mois e han an
nes s in he pine o es wi h he highes mois u e a
he op o an nes s. Ma e ial pH was simila in bo h
o es s ands and was lowe a he ops o an nes s
in compa ison wi h he bo oms and ims. The ca bon
con en was he highes a he ops o an nes s in bo h
o es s ands wi h g ea e diffe ences be ween sampling
loca ions in he pine o es . C:N a io was on a e age
highe in he pine o es wi h he highes alue a he
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Respi a ion o wood an nes ma e ial 1195
Fig. 1. Respi a ion o ma e ials pe g DW soil om sampling lo-
ca ions in he bi ch and pine o es s ( h ee-way ANOVA, F4,56
= 22.35, P<0.001). TOP e e s o he op o he an nes s,
BOTTOM o he base o he nes s, RIM o he pe iphe y o he
nes s, and HUMUS and SOIL o he humus laye and mine al soil,
espec i ely, as con ols. Columns wi h diffe en le e s a e s a-
is ically diffe en (one-way ANOVA, Tukey HSD pos hoc es ).
Means ±SEM a e shown.
Fig. 2. Respi a ion o ma e ials pe g ca bon om sampling lo-
ca ions in he bi ch and pine o es s ( h ee-way ANOVA, F4,56
= 1.42, P= 0.24). TOP e e s o he op o he an nes s, BOT-
TOM o he base o he nes s, RIM o he pe iphe y o he nes s,
and HUMUS and SOIL o he humus laye and mine al soil, e-
spec i ely, as con ols. Means ±SEM a e shown.
op o an nes s. In he bi ch o es , C:N a io did no
diffe be ween he sampling loca ions.
Discussion
In ag eemen wi h ou hypo heses, ma e ial espi a ion
pe g DW diffe ed be ween he sampling loca ions and,
also, be ween he wo o es s ands. This confi ms ea -
lie findings ha he same species can affec soil p op-
e ies diffe en ly in diffe en ecosys ems (F ouz e al.
2003; Holec & F ouz 2006). Respi a ion o he ma e-
ial om he op o an nes s in he bi ch o es was
significan ly highe han espi a ion in he o he pa s
o an nes s. Since li e decomposi ion is a complex
p ocess dependen on mois u e, empe a u e, o li e
quali y (Edwa ds e al. 1970; S eubing 1970; Paul &
Cla k 1996), his significan diffe ence could be mos
p obably explained by highe mois u e o he ma e ial
om he op o an nes s oge he wi h qui e low C:N
a io in he bi ch o es in compa ison wi h he pine o -
es . Howe e , in he pine o es he espi a ion o he
nes ma e ials om he diffe en pa s o an nes s was
simila . The eason o missing significan diffe ences
be ween mic osi es wi hin an nes s could be simila
mois u e in all he ma e ials and C:N a io in he ma-
e ial om he op wice as high as in he bo om and
im o an nes s. Despi e he low pH o he humus laye s
in bo h o es s ands, he humus laye espi a ion was
highe han in he bo oms and ims o an nes s, which
could be caused by highe mois u e and ca bon con en .
Appa en ly, he mois u e, C:N a io, and ca bon con-
en , sepa a ely o in combina ion, had an impo an
effec on ma e ial espi a ion in ou samples.
E en hough he empe a u e is conside ed o be
he mos impo an ac o influencing he a e o o -
ganic ma e u no e , i s o e all effec is complica ed
by he in e ac ion be ween empe a u e and mois u e
(Edwa ds e al. 1970). In addi ion o highe empe a-
u es eco ded inside an mounds (Coenen-S aß e al.
1980), an inc ease o mois u e was ound s imula ing de-
composi ion p ocesses o mound ma e ial (F ouz 1996;
Lenoi e al. 2001). Bu as mounds a e usually d ie
han he su ounding o es floo due o an cons uc -
ing ac i i ies (Sei e 1996; Lafleu e al. 2002), decom-
posi ion p ocesses can be lowe ed he e (Lenoi e al.
2001; Domisch e al. 2008). Ne e heless, decomposi ion
in an nes s in he pine o es was no dec eased due o
lowe mois u e con en , which sugges s ha he e a e
o he p ope ies affec ing his p ocess, e.g. ca bon a ail-
abili y. On he o he hand, ma e ials om an nes s in
he bi ch o es had on a e age highe mois u e han
in he pine o es , which means ha he li e decom-
posi ion was p omo ed he e p obably due o highe
mois u e con en . This esul is in acco dance wi h e-
sul s o Coenen-S aß e al. (1980), who ound highe
mois u e inside an nes s, and hus inc eased mic obial
ac i i y.
Ne e heless, he ma e ial espi a ion was affec ed
no only by mois u e, bu also by he C:N a io, which
was ound o be lowe in he bi ch o es and hus o be
ano he ac o s imula ing decomposi ion in his o es
s and. C:N a io a he op o an nes s in he pine o -
es was wice as high in compa ison wi h he bo om
and he im. Mound ma e ial a he op o an nes s
in he pine o es consis s p edominan ly o needles,
whe eas he bo om and he im consis o he mix-
u e o needles and mine al soil (K is iansen & Amelung
2001). Needles om coni e ous ees con ain esis an
cons i uen s such as cellulose and lignin, and o en ha e
la ge C:N a io (S eubing 1970). Hence, he C:N a io
a he op o an nes s in he pine o es was affec ed
by he high C:N a io o needles.
Howe e , since he a ailabili y o ca bon is e y
impo an o he espi a ion o o ganisms (Schulze e
al. 2000), we calcula ed he espi a ion pe g C and
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1196 V. Jilko a e al.
i showed diffe en esul s han he espi a ion pe g
DW. The espi a ion inc eased in an nes s, especially
in compa ison o he humus laye s. The mos impo an
p ope ies ha p obably affec ed he espi a ion we e
pH o he ma e ial and a ailabili y o ca bon. Highe
pH a o s he ac i i y o bac e ia mo e han ha o
ungi (Paul & Cla k 1996). Since bac e ia ha e highe
a e o me abolism, i can hus lead o highe CO2p o-
duc ion. In an nes s, an s accumula e as amoun s o
ood, mainly honeydew, which consis s o simple sug-
a s easily a ailable o mic oo ganisms. Mic oo ganisms
ha e enough ene gy o decomposi ion p ocesses and
ha also leads o highe CO2p oduc ion (i.e., p iming
effec ).
In conclusion, ou da a showed ha ma e ial de-
composi ion, and hus CO2efflux, in ou o es s ands
a e affec ed mainly by mois u e, ca bon con en and
a ailabili y, C:N a io, and pH. Al hough we examined
an mounds o he same species (Fo mica aquilonia)in
bo h o es s ands, ma e ial p ope ies o an nes s di -
e ed be ween he bi ch and he pine o es s. F om his
we can conclude ha nes ma e ial p ope ies a e a -
ec ed no only by he an species p esen and i s way
o li e, eeding and nes cons uc ion (Pe al 1978), bu
also by he o es s and cha ac e is ics.
Acknowledgemen s
The au ho s hank o he Uni e si y in Joensuu and he
Finnish Fo es Resea ch Ins i u e o p o iding he labo a-
o y equipmen . The s udy was financially suppo ed by he
g an MEYS CR (No LC06066).
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Accep ed Augus 16, 2013
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