Dehydrogenase Activity in a Litter Manipulation Experiment in Temperate Forest Soil

DOI: 10.2478/aslh-2013-0002 Ac a Sil . Lign. Hung., Vol. 9 (2013) 25–33 Dehyd ogenase Ac i i y in a Li e Manipula ion Expe imen in Tempe a e Fo es Soil Zsuzsa VERES* – Zsol KOTROCZÓ – Ko nél MAGYAROS – János A ila TÓTH – Béla TÓTHMÉRÉSZ Depa men o Ecology, Uni e si y o Deb ecen, Deb ecen, Hunga y Abs ac – Soil enzyme ac i i ies a e “senso s” o soil o ganic ma e (SOM) decomposi ion since hey in eg a e in o ma ion abou mic obial s a us and physico-chemical condi ion o soils. We measu ed dehyd ogenase enzyme ac i i y in a deciduous empe a e oak o es in Hunga y unde li e manipula ion ea men s. The Sík őkú De i us Inpu and Remo al T ea men s (DIRT) P ojec includes ea men s wi h doubling o lea li e and woody deb is inpu s as well as emo al o lea li e and enching o p e en oo inpu s. We hypo hesized ha inc eased de i al inpu s inc ease labile ca bon subs a es o soils and would inc ease enzyme ac i i ies pa icula ly ha o dehyd ogenase, which has been used as an indica o o soil mic obial ac i i y. We also hypo hesized ha enzyme ac i i ies would dec ease wi h de i us emo al plo s and dec ease labile ca bon inpu s o soil. A e en yea s o ea men s, li e emo al had a s onge e ec on soil dehyd ogenase ac i i y han did li e addi ions. These esul s showed ha in his o es ecosys em he changed li e p oduc ion a ec ed soil mic obial ac i i y: educed li e p oduc ion dec eased he soil dehyd ogenase ac i i y; inc eased li e p oduc ion had no signi ican e ec on he enzyme ac i i y. oak o es / dehyd ogenase ac i i y / Sík őkú P ojec / li e inpu / li e emo al / soil enzymes Ki ona – Az a a mennyiségének ha ása egy cse es- ölgyes e dő alajában a dehid ogenáz enzim ak i i ásá a. A alajenzimek a alajban lé ő sze es anyag (SOM) bomlásának "szenzo ai", mi el in o máció adnak a alaj mik obiológiai és izikai-kémiai állapo á ól. Egy magya o szági mé sékel ö i ölgye dőben mé ük a dehid ogenáz enzim ak i i ásá az a a mennyiségének csökken ésé el és nö elésé el. A Sík őkú DIRT P ojec ben (De i us Inpu and Remo al T ea men s) az alábbi kezeléseke alkalmaz uk: dupla mennyiségű le él a a , dupla mennyiségű ága a , alamin le él és gyöké meg onásos kezelések, ahol a gyöke eknek a pa cellák a ö énő benö ésé akadályozzuk meg. Az el é elez ük, hogy a megnö el a a inpu ha ásá a megnö ekszik a alajban a labilis, azaz a könnyen bon ha ó szén szubsz á ok mennyisége és az enzimek ak i i ása. Különösen a dehid ogenáz enzim ak i i ásának ál ozásá á uk, ami az egyik legál alánosabban használ mu a ó a alaj mik obiális ak i i ásának mé ésé e. To ábbá az el é elez ük, hogy a alaj enzim ak i i ása csökkenni og az a a meg onásos kezelésekben, a labilis szén szubsz á ok mennyiségének csökkenésé el. E edményeink az mu a ják, hogy íz é el el é el az a a p odukció csökkenése ebben az e dei ökoszisz émában a alaj dehid ogenáz enzim ak i i ásának csökkenésé okoz a, ugyanakko az a a p odukció nö ekedése nem okozo szigni ikáns ál ozás az enzim ak i i ásában ölgye dő / dehid ogenáz ak i i ás / Sík őkú P ojec / a a mennyiség megduplázása / a a meg onás / alajenzimek * Co esponding au ho : e [email protected]; H-4032 DEBRECEN, Egye em é 1. Unau hen ica ed Download Da e | 1/21/15 1:14 PM Ve es, Zs. e al. Ac a Sil . Lign. Hung. 9, 2013 26 1 INTRODUCTION Soil enzyme ac i i ies a e “senso s” o soil o ganic ma e (SOM) decomposi ion since hey in eg a e in o ma ion abou mic obial s a us and physico-chemical condi ion o soils (Aon – Colane i 2001, Baum e al. 2003). All soils con ain a g oup o enzymes ha de e mine soil me abolic p ocesses (McLa en 1975) which, in u n, depend on i s physical, chemical, mic obiological and biochemical p ope ies (Makoi and Ndakidemi 2008). Enzyma ic p ocesses a e closely ela ed o soil quali y, hey pa icipa e in he p ocessing o una ailable o ms o nu ien s eadily assimila ed by plan s (Sinsabaugh e al. 1994). Soil dehyd ogenase enzymes (DHA) (EC 1.1.1.1) a e one o he main componen s o soil enzyma ic ac i i ies pa icipa ing in and assu ing he co ec sequence o all he biochemical ou es in soil biogeochemical cycles (Ladd 1985). De e mina ion o DHA in soil gi es la ge amoun o in o ma ion abou biological cha ac e is ic o he soil (Wolińska – S ępniewska 2012). Dehyd ogenase enzyme is o en used as a measu e o any dis up ion caused by pes icides, ace elemen s o managemen p ac ices o he soil, as well as a di ec measu e o soil mic obial ac i i y (Chend ayan e al. 1979; T e o s e al. 1982; McCa hy e al. 1994). Soil DHA is conside ed o exis in soils as in eg al pa s o in ac cells (Taylo e al. 2002). Dehyd ogenase enzyme is known o oxidize soil o ganic ma e by ans e ing p o ons and elec ons om subs a es o accep o s (Makoi – Ndakidemi 2008). These p ocesses a e pa o espi a ion pa hways o soil mic o-o ganisms and a e closely ela ed o he ype o soil and soil ai -wa e condi ions (Kandele e al. 1996). Se e al en i onmen al ac o s, including soil mois u e, oxygen a ailabili y, oxida ion educ ion po en ial, pH, o ganic ma e con en , dep h o he soil p o ile, empe a u e, season o he yea , hea y me al con amina ion and soil e iliza ion o pes icide use can a ec signi ican ly DHA in he soil en i onmen (Dkha – Mish a 1983; Ba uah – Mish a 1984; Sinsabaugh e al. 2008; Xiang e al. 2008; Wolińska – S ępniewska 2012; Kuma e al. 2013). We measu ed DHA in a deciduous empe a e oak o es in Hunga y unde li e manipula ion ea men s. Ou esea ch in he Sík őkú DIRT P ojec (SIK) cons i u es an impo an pa o an in e na ional long e m p ojec which in ol es i e expe imen al si es in he USA (H.J. And ews Expe imen al Fo es , Bousson Expe imen al Fo es , Ha a d Fo es , Uni e si y o Michigan Biological S a ion, San a Ri a) (Nadelho e e al. 2004) and one in Ge many (Uni e si ä Bay eu h BITÖK). We hypo hesized ha inc eased de i al inpu s would p o ide inc eased labile ca bon subs a es o soils and inc ease dehyd ogenase ac i i ies, which is used as indica o o soil mic obial ac i i y (Thalmann 1968a; Chend ayan e al. 1979; Nannipie i e al. 2002). We also hypo hesized ha dehyd ogenase ac i i ies would dec ease in de i us emo al plo s wi h dec eases in labile ca bon inpu s o soil. 2 METHODS 2.1 A ea and p ojec desc ip ions The Sík őkú P ojec was es ablished in 1972 o he long- e m s udy o o es ecosys ems. I is loca ed in he sou h pa o he Bükk Moun ains in No h-Eas e n Hunga y N 47°55’ E 20°46’ a 325 m al i ude. This o es has been p o ec ed since 1976, and i is pa o he Bükk Na ional Pa k. Mean annual empe a u e (MAT) is 10°C and Mean Annual P ecipi a ion (MAP) is 550 mm (An al e al. 1997). Soil pH anged be ween 4.85 and 5.50 depending on he plo s (Tó h e al. 2007). Bo h soil pH and humus con en was lowe in de i us emo al ea men s. Humus con en anged be ween 2.66% and 3.14% (Va ga e al. 2008). In he con ol plo s he ca bon con en was 5.19% (0–5 cm), and 3.25% (5–15 cm) (Tó h e al. 2011). Acco ding o he FAO Soil Classi ica ion, he soil is a Cambisols. This Unau hen ica ed Download Da e | 1/21/15 1:14 PM Dehyd ogenase ac i i y in a li e manipula ion expe imen Ac a Sil . Lign. Hung. 9, 2013 27 o es is a semi-na u al s and (Que ce um pe aeae-ce is communi y) wi hou o es managemen ac i i y (Jakucs 1985; Ko oczó e al. 2012). The Sík őkú P ojec is a membe si e o he Hunga ian LTER Ne wo k (Long Te m Ecological Resea ch) and he ILTER Ne wo k (In e na ional Long Te m Ecological Resea ch) om 1995 (Ko ács-Láng e al. 2000). Six ea men s we e es ablished a he Sík őkú DIRT expe imen al si e in he au umn o 2000 each in h ee eplica es on 7  7 m (49 m2) plo s. De i us was no manipula ed in he Con ol plo s (CO). The e we e no mal li e inpu s in he CO plo s. The a e age lea -li e p oduc ion was 3547 kg ha–1yea –1 be ween 2003 and 2010 (Ko oczó e al. 2012). The e we e wo ypes o de i us addi ions: double he no mal amoun o lea li e was applied o he Double Li e (DL) plo s by adding lea li e emo ed om NL plo s; while in he Double Wood plo s (DW) he amoun o wood de i us (b anches, wigs and ba k) was doubled. Annual wood li e amoun was measu ed by boxes placed o he si e and i s double amoun was applied in he case o e e y DW plo s (a e age 17 kg yea –1). In h ee ea men s o de i us emo al we e also applied: abo eg ound de i us emo ed by ake in he No Li e plo s (NL), li ing oo s se e ed by enching in he No Roo s plo s (NR), and bo h NL and NR ea men s in No Inpu s plo s (NI). The NR and NI plo s we e enched a ound 40 cm wide and 100 cm deep. The soil dug ou was placed ou side he plo . Roo -p oo Del a MS 500 PE oil was pu in he enches, which was 0.6 mm hick and 1 m wide. Then he enches we e illed wi h soil. Plan s we e clea ed o elimina e oo p oduc ion (bushes we e cu ou a he es ablishmen ). Weeds we e also con olled by Medalon (agen : 480 g l–1 gli osa e- ammonium) and d y plan esidues we e aked. The de ailed desc ip ion o he ea men s is in Nadelho e e al. (2004), Sulzman e al. (2005), Ko oczó e al. (2008). These p ocesses we e eplayed e e y yea . 2.2 Soil sampling and measu ing o dehyd ogenase enzyme ac i i y Soil samples we e collec ed 11 imes om Feb ua y 2010 o No embe 2012. Fi e co es we e aken om each plo 15 cm dep h wi h a 2 cm diame e Oak ield soil co e (Oak ield Appa a us Company, USA). Th ee analy ical eplica es pe sample pe assay we e used. The samples we e homogenized and s o ed o one week a 4°C. DHA was de e mined using he educ ion o 2.3.5- iphenyl e azolium chlo ide (TTC) me hod (Thalmann 1968b). Samples o 2 g ield-mois soils we e mixed wi h 2 ml 1.5% TTC ho oughly in o es ubes. The samples we e mixed on a o ex and incuba ed a 30°C. The con ol con ains only 2 ml T is bu e (wi hou TTC). A e 24 h, he iphenyl o mazan, a p oduc om he educ ion o TTC, was ex ac ed by adding 10 ml e hanol o each ube and shaken o 1 min. Samples we e u he incuba ed a oom empe a u e o 2 h in he da k (shaking he ubes a in e als). The soil suspensions (12 ml) we e hen il e ed and he op ical densi y o he clea supe na an was measu ed agains he blank (e hanol) a 546 nm ( ed colo ) a Genesys 10 spec opho ome e . A s anda d cu e was plo ed using a ange o iphenyl o mazan (TPF) (Reanal, Budapes , Hunga y) concen a ions be ween 0 and 40 µg TPF ml–1. DHA was exp essed as µg TPF g–1 d y soil 24 h–1. Soil mois u e con en was measu ed by TDR 300 (Time Domain Re lec ome e ) ins umen in he ield in olume ic wa e con en pe cen ( wc%). The e we e wo measu emen s a each plo a he ime o soil sampling o de e mine DHA. Soil empe a u e was measu ed by an ONSET, S owAway TidbiT- ype da a-logge (Onse Compu e Co po a ion, USA), in he middle o each plo a 10 cm dep h. Da a-logge s we e p og ammed o measu e soil empe a u e e e y hou ; daily mean o he soil empe a u e alues was used in he pape . Unau hen ica ed Download Da e | 1/21/15 1:14 PM Ve es, Zs. e al. Ac a Sil . Lign. Hung. 9, 2013 28 2.3 S a is ical analyzes We used one-way ANOVA o de e mine signi ican ea men e ec s; signi ican mean di e ences we e e alua ed a he le el o p=0.05 using Tukey’s es . 3 RESULTS AND DISCUSSION We s a ed o measu e soil DHA in 2010, en yea s a e he es ablishmen o ea men s. In gene al, ac i i ies we e lowe in emo al ea men s (NL, NR, NI) han he con ol (CO) o addi ion ea men s (DL, DW). Ou da a show ha he dehyd ogenase ac i i y is ai ly s able in he con ol plo s compa ed o he ea ed plo s (Figu e 1). The e is a empo al end in dehyd ogenase ac i i y; he e ec o de i us emo al is inc eased. Figu e 1. Soil dehyd ogenase enzyme ac i i y among ea men s be ween 2010 and 2012 (µg TPF g–1 d y soil 24h–1± SE). Di e en le e s deno e signi ican ea men di e ences wi hin each yea (p<0.05, ANOVA and Tukey's es ). DHA we e signi ican lowe in NR han in DL ea men s in 2010 (p=0.029). In 2011 he e we e highe signi ican di e ences in dehyd ogenase ac i i y among ea men s, enzyme ac i i y was highe in DL and DW ea men s han in all emo al plo s (p<0.001). In 2012 li e emo al ea men s di e ed signi ican ly no only o DL and DW ea men s bu also con ols (p<0.001) (Figu e 1). The enzyme ac i i ies we e highe in DL and DW ea men s han con ols, bu no signi ican ly. In con as o ou ini ial hypo hesis, dehyd ogenase ac i i ies did no inc ease signi ican ly wi h ei he lea li e o wood addi ions. This is gene ally in ag eemen wi h ou ea lie esul s o li e decomposi ion (Feke e e al. 2007), soil espi a ion (Ko oczó e al. 2008), a ylsulpha ase and saccha ase ac i i ies (Feke e e al. 2011). A he o he DIRT si e in H.J. And ews Expe imen al Fo es (HJA) in O egon, B an e al. (2006a) showed ha , soil om he doubled wood ea men had a highe ungal:bac e ial a io, and soil om he no inpu s ea men had a lowe ungal:bac e ial a io, han he con ol soil. These changes we e he esul o al e a ion in he size and composi ion o he mic obial Unau hen ica ed Download Da e | 1/21/15 1:14 PM Dehyd ogenase ac i i y in a li e manipula ion expe imen Ac a Sil . Lign. Hung. 9, 2013 29 communi y. B an e al. (2006b) epo ed mic obial biomass alues a h ee DIRT si es: HJA, SIK and BOU (Bousson Expe imen al Fo es in Pennsyl ania). They ound signi ican di e ences in biomass be ween ea men s a SIK. A ha si e DW plo s had a la ge biomass han he NR plo s and sugges i e e idence ha he DW plo s had a la ge biomass han he NI plo s. Błońska e al. (2013) epo ha , DHA is inc easing wi h he di e si y o plan communi ies; his sugges ha he e is a ela ionship be ween he composi ion o soil o ganic ma e and mic obial enzyma ic ac i i y. Mama ha e al. (2001) s udying soil om ed sandy loam a eas show ha , he DHA is highe in soil co e ed by plan s and in he hizosphe e han non- hizosphe e. Table 1. Mean alues and s anda d e o s o dehyd ogenase ac i i y, soil mois u e con en and soil empe a u e in e ms o he ea men s. (Soil mois u e and empe a u e alues a e he mean alues o he days o he measu emen s). Signi ican di e ences in dehyd ogenase ac i i y be ween ea men s showed in Figu e 1 (p<0.05, ANOVA and Tukey's es ). T ea men s Dehyd ogenase ac i i y (µg TPF g–1 d y soil 24h–1± SE) Soil mois u e ( wc% ± SE) Soil empe a u e (°C) 2010 NL 0.43±0.12 34.07±1.95 8.29±4.09 NR 0.32±0.12 41.00±2.01 9.63±4.20 NI 0.41±0.10 38.36±2.02 9.46±4.26 CO 0.62±0.12 34.84±2.63 8.92±3.86 DL 0.93±0.17 34.38±2.56 9.28±3.64 DW 0.65±0.17 35.07±2.99 9.60±4.14 p=0.029 p>0.05 p>0.05 2011 NL 0.20±0.08 18.82±3.39 17.69±3.21 NR 0.29±0.11 25.89±4.84 17.93±3.22 NI 0.22±0.08 21.68±4.44 18.02±3.30 CO 0.73±0.13 20.88±4.44 17.86±3.20 DL 0.98±0.20 21.87±4.41 17.13±2.96 DW 1.17±0.23 21.30±4.62 18.07±3.27 p<0.001 p>0.05 p>0.05 2012 NL 0.14±0.03 19.93±3.22 12.50±5.68 NR 0.17±0.03 20.66±3.59 12.46±5.75 NI 0.09±0.03 21.32±3.87 12.52±5.76 CO 0.60±0.06 16.18±2.51 12.47±5.76 DL 0.78±0.10 17.36±2.67 12.87±5.34 DW 0.68±0.10 18.26±3.12 12.58±5.75 p<0.001 p>0.05 p>0.05 We measu ed he soil mois u e and soil empe a u e a he days when soil sampling was made o de e mine DHA. We did no ind signi ican ea men di e ences wi hin each yea in soil mois u e and in soil empe a u e (Table 1). The soil mois u e alues we e he highes in 2010 (Figu e 2); his yea was ex emely ainy. Con a ily, yea s 2011 and 2012 we e d y and wa m; ins ead o ha he DHA showed same alues in he CO plo s han in 2010. In he Unau hen ica ed Download Da e | 1/21/15 1:14 PM Ve es, Zs. e al. Ac a Sil . Lign. Hung. 9, 2013 30 emo al plo s DHA dec eased in 2011 and 2012. P obably he highe soil mois u e caused he highe DHA in 2010. Howe e , we did no ind co ela ion be ween soil mois u e and DHA in yea s 2010, 2011 and 2012. Kuma e al. (2013) showed signi ican co ela ion be ween DHA and soil mois u e. O he au ho s also a ibu ed he inc ease in mic obial ac i i y in o es soil o highe soil mois u e (Gö es e al. 1998). Feke e e al. (2011) also ound co ela ion be ween soil mois u e and enzyme ac i i ies (a ylsulpha ase and saccha ase) in Sík őkú be ween 2004 and 2006. Ou esul s sugges ha , changes in soil o ganic ma e con en had highe e ec on DHA han soil mois u e. In o he s udies also ound ha mo e o ganic ma e main ained la ge and mo e ac i e mic obial biomass and highe DHA (Chodak e al. 2010; Bonanomi e al. 2011). Figu e 2. Soil mois u e con en ( wc%) among ea men s be ween 2010 and 2012. A e en yea s o ea men s, li e emo al had a s onge e ec h ough ime on soil dehyd ogenase ac i i ies han did inc ease li e inpu s. Ec omyco hizal ungi associa ed wi h oo s o ees in deciduous o es s p oduce enzymes and hus can inc ease he su ounding soil enzyme ac i i ies (Colpae – an Lae e 2006; Smi h – Read 2008). These ungi a e esponsible o he mobiliza ion o essen ial plan nu ien s by decomposing li e and soil o ganic ma e ials (Cou y e al. 2006). Decomposi ion o se e ed oo s (NR and NI) would esul in he disappea ance o myco hizal ungi and hizosphe e mic obes, causing a conside able educ ion in mic obial and enzyme ac i i ies (Feke e e al. 2011). 4 CONCLUSION Ou indings showed ha he changes o li e p oduc ion signi ican ly a ec ed soil mic obial ac i i y, and by ex ension gene al nu ien cycling and SOM dynamics in his o es ecosys em. When he li e p oduc ion dec eased, soil dehyd ogenase ac i i y also dec eased. Inc eased li e p oduc ion caused no signi ican inc ease in enzyme ac i i y. A e en yea s o ea men s, loss o inpu s had a g ea e impac han inc eased inpu s. Unau hen ica ed Download Da e | 1/21/15 1:14 PM Dehyd ogenase ac i i y in a li e manipula ion expe imen Ac a Sil . Lign. Hung. 9, 2013 31 Acknowledgmen s: We hank Csabáné Koncz o long hou s o ield and labo a o y wo k. Special hanks a e due o D . Ka e Laj ha and B uce Caldwell (O egon S a e Uni e si y, Co allis, USA) and D . K is in Vande bil (Uni e si y o New Mexico, Albuque que, USA) o hei subs an ial con ibu ion in es ablishing he expe imen . This esea ch was suppo ed by he Eu opean Union and he S a e o Hunga y, co- inanced by he Eu opean Social Fund in he amewo k o TÁMOP 4.2.4. A/2-11-1-2012-0001 ‘Na ional Excellence P og am’. The wo k was suppo ed by TÁMOP 4.2.1./B-09/1/KONV-2010-0007, TÁMOP-4.2.2_B-10_1- 2010-0024 and TÁMOP-4.2.2.C-11/1/KONV-2012-0010 p ojec s. The TÁMOP p ojec s a e implemen ed h ough he New Hunga y De elopmen Plan, co- inanced by he Eu opean Social Fund and he Eu opean Regional De elopmen Fund. REFERENCES ANTAL, E. – BERKI, I. – JUSTYÁK, J. – KISS, GY. – TARR, K. – VIG, P. (1997): A sík őkú i e dő á sulás hő- és ízház a ási iszonyainak izsgála a az e dőpusz ulás és az éghajla ál ozás ük ében, Deb ecen. pp 83. (in Hunga ian). AON, M. A. – COLANER, A. C. (2001): Tempo al and spa ial e olu ion o enzyma ic ac i i ies and physico-chemical p ope ies in an ag icul u al soil. 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