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  • Arnstadt, Tobias  (11)
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  • 1
    Language: English
    In: Forest Ecology and Management, 15 December 2016, Vol.382, pp.129-142
    Description: In forest ecosystems, deadwood is an important component that provides habitat and contributes to nutrient cycles, as well as to carbon and water storage. The change of wood constituents, nutrients and microbial species richness in the field over the whole time of decomposition has only rarely been studied, in particular not in relation to oxidative enzyme activities (mediating lignin degradation) and different forest management regimes. To describe wood decomposition, we selected coarse woody debris (CWD) in form of 197 logs of , and in forests with different management regimes across three regions in Germany. They were sampled and analyzed for wood density, water content, wood constituents (Klason and acid-soluble lignin, organic extractives, water-soluble lignin fragments), carbon, nitrogen and metals (Al, Ca, Cu, K, Mg, Mn and Zn). Furthermore, the activities of oxidative enzymes like laccase, manganese peroxidase, and general peroxidase were measured. Since filamentous fungi (Basidiomycota, Ascomycota) are the major biological agents of wood decomposition, fungal species richness based on sporocarps and molecular fingerprints was recorded. Higher forest management intensity had a negative effect on deadwood volume and in consequence on fungal species richness (sporocarps), but hardly to other analyzed variables. Furthermore, there were significant differences between the tree species for the concentrations of wood constituents and most nutrients as well as the activities of oxidative enzymes, although their course during decomposition was mostly similar among the tree species. We found that molecular species richness increased with the period of decomposition in contrast to the number of fruiting species, which was highest in the intermediate stage of decomposition. Both types of species richness increased with increasing volume of the CWD logs. Regarding the entire period of decomposition, white-rot fungi (WRF), based on identification of sporocarps, were the most abundant group of wood-decaying fungi in all three tree species. This corresponds well with the overall presence of laccase and peroxidases and the concomitant substantial loss of lignin, which points to the importance of these enzymes in deadwood decomposition. We found a continuous decomposition and decline of volume-related concentrations in wood constituents and nutrients with time of decomposition. Contrary to volume-related concentrations, the concentrations related to dry mass frequently increased.
    Keywords: Saproxylic Fungi ; Oxidative Enzyme ; Nutrients ; Lignin ; Forest Management Intensity ; Coarse Woody Debris ; Forestry ; Biology
    ISSN: 0378-1127
    E-ISSN: 1872-7042
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  • 2
    Language: English
    In: Forest Ecology and Management, 01 May 2017, Vol.391, pp.86-95
    Description: Deadwood decay is an important ecosystem process in forest ecosystems, but the relative contribution of specific wood properties of tree species, activities of wood-degrading enzymes, and decomposer communities such as fungi and insects is unclear. We ask whether wood properties, in particular differences between angiosperms and gymnosperms, and organismic diversity of colonizers contribute to wood decomposition. To test this, we exposed deadwood logs of 13 tree species, covering four gymnosperms and nine angiosperm species, in 30 plots under different forest management in three regions in Germany. After a decomposition time of 6.5 years and showed the highest decay rates. We found a positive correlation of decay rate with enzyme activities, chemical wood properties (S, K concentration) and organismic diversity, while, heartwood character, lignin content, extractive concentration and phenol content were negatively correlated with decay rate across all 13 tree species. By applying a multi-model inference approach we found that the activity of the wood-degrading enzymes laccase and endocellulase, beetle diversity, heartwood presence, wood ray height and fungal diversity were the most important predictor variables for wood decay. Although we were not able to identify direct cause and effect relations by our approach, we conclude that enzyme activity and organismic diversity are the main drivers of wood decay rate, which greatly differed among tree species. Maintaining high tree species diversity will therefore result in high structural deadwood diversity in terms of decay rate and decay stage.
    Keywords: Wood Decomposition ; Ecosystem Function ; Saproxylic Beetles ; Biodiversity Exploratories ; Deadwood Experiment ; Forestry ; Biology
    ISSN: 0378-1127
    E-ISSN: 1872-7042
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  • 3
    In: Scientific Reports, 2015, Vol.5
    Description: Deadwood is an important biodiversity hotspot in forest ecosystems. While saproxylic insects and wood-inhabiting fungi have been studied extensively, little is known about deadwood-inhabiting bacteria. The study we present is among the first to compare bacterial diversity and community structure of deadwood under field conditions. We therefore compared deadwood logs of two temperate forest tree species Fagus sylvatica and Picea abies using 16S rDNA pyrosequencing to identify changes in bacterial diversity and community structure at different stages of decay in forest plots under different management regimes. Alphaproteobacteria, Acidobacteria and Actinobacteria were the dominant taxonomic groups in both tree species. There were no differences in bacterial OTU richness between deadwood of Fagus sylvatica and Picea abies. Bacteria from the order Rhizobiales became more abundant during the intermediate and advanced stages of decay, accounting for up to 25% of the entire bacterial community in such logs. The most dominant OTU was taxonomically assigned to the genus Methylovirgula, which was recently described in an experiment of Fagus sylvatica woodblocks. Besides tree species we were able to demonstrate that deadwood physico-chemical properties, in particular remaining mass, relative wood moisture, pH, and C/N ratio serve as drivers of community composition of deadwood-inhabiting bacteria.
    Keywords: Species ; Fungi ; Decay ; Carbon/Nitrogen Ratio ; Herbivores ; Community Structure ; Biodiversity ; Rrna 16s ; Rainforests ; Physicochemical Properties ; Bacteria ; Community Composition ; Ph Effects;
    ISSN: 20452322
    E-ISSN: 20452322
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  • 4
    Language: English
    In: European Journal of Forest Research, 2016, Vol.135(1), pp.109-124
    Description: Lignin and its degradation, particularly in forest ecosystems, play a major role in the global carbon cycle. Filamentous fungi equipped with extracellular oxidoreductases (oxidative enzymes), i.e., laccase, manganese-dependent peroxidases and several other peroxidases, are the key players in the bioconversion of lignin. In particular, for coarse woody debris (CWD), this process is poorly understood and the activities of laccase and peroxidases have never been studied on a large field scale. We investigated the activities of these enzymes in 701 samples of Fagus sylvatica , Picea abies and Pinus sylvestris CWD across three regions in Germany and analyzed their dependence on pH, water content, wood density, total lignin, organic extractives, metals, water-soluble lignin fragments and fungal species richness. Respective enzyme activities were present in 79 % of all samples, and the activities were highly variable and more frequent in F. sylvatica than in coniferous wood. Logistic regressions and correlations between enzyme activities and the variables revealed that the fungal community structure and the amount of water-soluble lignin fragments are most important determinants, and that the prevalent acidic pH in CWD is suitable to facilitate laccase and manganese peroxidase activities. Concentrations of metals (manganese, copper, iron) were sufficient to ensure synthesis and functioning of relevant enzymes. Based on this large field study, we conclude that laccase and peroxidases in CWD are highly relevant for lignin degradation, but the variable pattern of their secretion is the result of a complex array of wood parameters and the fungal community structure, which could only partly be resolved.
    Keywords: Laccase ; Manganese peroxidase ; General peroxidase ; Dead wood ; Lignin ; Wood rot
    ISSN: 1612-4669
    E-ISSN: 1612-4677
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  • 5
    Language: English
    In: Fungal Diversity, 2016, Vol.77(1), pp.367-379
    Description: Fungi play vital roles in the decomposition of deadwood due to their secretion of various enzymes that break down plant cell-wall complexes. The compositions of wood-inhabiting fungal (WIF) communities change over the course of the decomposition process as the remaining mass of wood decreases and both abiotic and biotic conditions of the wood significantly change. It is currently not resolved which substrate-related factors govern these changes in WIF communities and whether such changes influence the deadwood decomposition rate. Here we report a study on fungal richness and community structure in deadwood of Norway spruce and European beech in temperate forest ecosystems using 454 pyrosequencing. Our aims were to disentangle the factors that correspond to WIF community composition and to investigate the links between fungal richness, taxonomically-resolved fungal identity, and microbial-mediated ecosystem functions and processes by analyzing physico-chemical wood properties, lignin-modifying enzyme activities and wood decomposition rates. Unlike fungal richness, we found significant differences in community structure between deadwood of different tree species. The composition of WIF communities was related to the physico-chemical properties of the deadwood substrates. Decomposition rates and the activities of lignin-modifying enzymes were controlled by the succession of the fungal communities and competition scenarios rather than fungal OTU richness. Our results provide further insights into links between fungal community structure and microbial-mediated ecosystem functions and processes.
    Keywords: Ecosystem processes ; Enzyme activity ; Wood decay ; Physico-chemical wood properties ; Ascomycetes ; Basidiomycetes ; Pyrosequencing
    ISSN: 1560-2745
    E-ISSN: 1878-9129
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  • 6
    Language: English
    In: Fungal Ecology, August 2016, Vol.22, pp.98-105
    Description: Wood-inhabiting fungi are major agents of wood decomposition. However, it is unclear which factors determine their distribution and enzyme production. Many studies that have addressed this issue suffer from a lack of geographic extent. Here, we investigate the fungal community structure of 117 logs in relation to wood physico-chemical properties and secreted ligninolytic enzymes, across three distinct geographical regions of Germany. Our results revealed that fungal community structure was similar across different regions, but was nevertheless variable in all regions. The relationships between fungal community structure, wood physico-chemical properties and enzyme activities were not consistent across different regions. However, we identified that the wood physico-chemical properties (i.e. decay class, remaining mass, density, extractives, total lignin and pH) were the most important factors associated with the fungal community structure in all three regions. In contrast, the wood physico-chemical properties and the fungal community structure did not sufficiently explain variation in the detected enzymatic activities. Thus, we assume that interspecific interactions and recently described priority effects play more important roles in the production of lignin modifying enzymes.
    Keywords: Ecosystem Processes ; Enzyme Activity ; Wood Decay ; Wood Physico-Chemical Properties ; Fungal Community Structure ; Botany
    ISSN: 1754-5048
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  • 7
  • 8
    In: Environmental Microbiology, May 2018, Vol.20(5), pp.1693-1710
    Description: Nitrogen deposition can strongly affect biodiversity, but its specific effects on terrestrial microbial communities and their roles for ecosystem functions and processes are still unclear. Here, we investigated the impacts of N deposition on wood‐inhabiting fungi (WIF) and their related ecological functions and processes in a highly N‐limited deadwood habitat. Based on high‐throughput sequencing, enzymatic activity assay and measurements of wood decomposition rates, we show that N addition has no significant effect on the overall WIF community composition or on related ecosystem functions and processes in this habitat. Nevertheless, we detected several switches in presence/absence (gain/loss) of wood‐inhabiting fungal OTUs due to the effect of N addition. The responses of WIF differed from previous studies carried out with fungi living in soil and leaf‐litter, which represent less N‐limited fungal habitats. Our results suggest that adaptation at different levels of organization and functional redundancy may explain this buffered response and the resistant microbial‐mediated ecosystem function and processes against N deposition in highly N‐limited habitats.
    Keywords: Biodiversity -- Environmental Aspects ; Biodiversity -- Analysis ; Ecosystems -- Environmental Aspects ; Ecosystems -- Analysis;
    ISSN: 1462-2912
    E-ISSN: 1462-2920
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  • 9
    Language: English
    In: Frontiers in Microbiology, 01 September 2018, Vol.9
    Description: Despite the important role of wood-inhabiting fungi (WIF) in deadwood decomposition, our knowledge of the factors shaping the dynamics of their species richness and community composition is scarce. This is due to limitations regarding the resolution of classical methods used for characterizing...
    Keywords: Next Generation Sequencing ; Microbial Ecology ; Belongdead ; Wood-Physicochemical Properties ; Fungal Richness ; Fungal Community Composition ; Biology
    E-ISSN: 1664-302X
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  • 10
    Language: German
    Description: In Waldökosystemen ist Totholz von zentraler Bedeutung, indem es zahlreichen Organismen einen Lebensraum bietet oder als Substrat dient, Bestandteil des Kohlenstoff- und Nährstoffkreislaufs ist sowie als ein wichtiges strukturelles Element fungiert. Für seine Zersetzung ist die Überwindung der Ligninbarriere von besonderer Bedeutung. Dazu sind lediglich saprobionte Pilze aus den Phyla der Basidiomycota und Ascomycota in der Lage, die verschiedene Strategien – die Fäuletypen – entwickelt haben, um Lignin abzubauen oder zu modifizieren und somit Zugang zu den vom Lignin inkrustierten Polysachariden (Zellulose und Hemizellulosen) zu erhalten. Eine besondere Rolle spielen dabei Weißfäulepilze, die mit ihren extrazellulären oxidativen Enzymen, wie Laccasen und verschiedenen Peroxidasen, Lignin komplett bis zum Kohlendioxid (CO2) mineralisieren. Trotz der Bedeutung des Ligninabbaus für die Totholzzersetzung sind extrazelluläre oxidative Enzyme im natürlichen Totholz kaum erforscht. Ziel dieser Arbeit war es, die Rolle der oxidativen Enzyme für die Totholzzersetzung unter Realbedingungen zu verifizieren, ihre räumlichen und zeitlichen Muster zu beschreiben und ihre Abhängigkeiten von verschiedenen Totholzvariablen sowie der pilzlichen Artengemeinschaft in und auf Totholz zu ermitteln. Weiter wurde die Veränderung der Totholzvariablen über den Zersetzungsprozess für unterschiedliche Baumarten vergleichend beschrieben und der Einfluss der Waldbewirtschaftung auf den Prozess untersucht. Dazu wurden 197 natürliche Totholzstämme (coarse woody debris, CWD) von Fagus sylvatica (Rotbuche), Picea abies (Gemeine Fichte) und Pinus sylvestris (Gemeine Kiefer) in unterschiedlich stark bewirtschafteten Wäldern in Deutschland untersucht. Insgesamt wurden 735 Proben genommen und darin die Aktivität von Laccase (Lacc), Genereller Peroxidase (GenP) und Mangan-Peroxidase (MnP) gemessen. Weiterhin wurden Variablen wie Dichte, Wassergehalt, pH-Wert, wasserlösliche Ligninfragmente, die Gehalte an Lignin und Extraktiven sowie an Nährstoffen und Metallen (N, Al, Ca, Cu, K, Mg, Mn und Zn) ermittelt. Die pilzliche Artengemeinschaft wurde anhand genetischer Fingerprints (F-ARISA) und mittels Fruchtkörperkartierung erfasst. In 79 % der untersuchten Totholzproben wurden oxidative Enzymaktivitäten festgestellt. Sie waren hoch variabel über den Zersetzungsverlauf sowie in Bezug auf die Probenahmepositionen innerhalb der einzelnen Stämme. Generell waren die Aktivitäten im F.-sylvatica-Totholz höher als im Koniferentotholz. Lineare und logistische Modelle zeigten, dass die pilzliche Artengemeinschaft, gefollgt von den wasserlöslichen Ligninfragmenten, die wichtigste Einflussgröße hinsichtlich der oxidativen Enzyme war. Ein saurer pH-Wert unterstützte die Funktion von Lacc und MnP; Mangan, Eisen und Kupfer waren in ausreichenden Konzentrationen vorhanden, um die Funktion und Bildung der Enzyme zu gewährleisten. Die holzabbauenden Pilze erwiesen sich als optimal an das niedrige Stickstoffangebot im Totholz angepasst, sodass ein erhöhter Stickstoffeintrag über zwei Jahre die oxidativen Enzymaktivitäten nicht weiter beeinflusste. Der pH-Wert sowie die Gehalte an Lignin, Extraktiven und Nährstoffen waren im Vergleich der drei Baumarten signifikant verschieden, obwohl die zeitlichen Veränderungen der Variablen über den Zersetzungsprozess vergleichbar waren. Die Anzahl operativer taxonomischer Einheiten (OTUs ~ molekulare Artenzahl) nahm im Verlauf der Holzzersetzung zu, während die Zahl fruktifizierender Arten für mittlere Zersetzungsgrade am höchsten war. Beide Artenzahlen nahmen zusammen mit dem Stammvolumen zu. Die Weißfäulepilze dominierten über den gesamten Zersetzungsprozess die fruchtkörperbasierte Artenzahl aller drei Baumarten, was mit dem Vorhandensein oxidativer Enzymaktivitäten einhergeht. Generell nahmen der massebezogene Gehalt des Lignins, der Extraktive und der Nährstoffe über die Zersetzung zu, während der volumenbezogene Gehalt abnahm. Der pH-Wert im Holz aller drei Baumarten sank kontinuierlich im Verlauf der Zersetzung. Eine Erhöhung der Waldbewirtschaftungsintensität hatte einen negativen Effekt auf das Stammvolumen und darüber vermittelt auf die Zahl fruktifizierender Pilzarten, jedoch kaum auf andere untersuchte Totholzvariablen. Aufgrund des häufigen Vorkommens von Weißfäulepilzen, der gleichzeitigen Präsenz oxidativer Enzymaktivitäten und des substanziellen Ligninabbaus kann auf eine fundamentale Bedeutung von Laccasen und Peroxidasen für die Zersetzung des Totholzes geschlossen werden. Nicht zuletzt die charakteristische Molekularmassenverteilung der wasserlöslichen Ligninfragmente deutete darauf hin, dass die Mn-oxidierenden Peroxidasen (MnPs) die dominierenden oxidativen Enzyme des Ligninabbaus sind. Das hoch variable Muster der oxidativen Enzymaktivitäten ist jedoch das Resultat eines komplexen Zusammenspiels der Holzeigenschaften und der pilzlichen Artengemeinschaft. Die dabei bestehenden funktionellen Abhängigkeiten müssen weiter im Detail in zukünftigen Studien analysiert und aufgeklärt werden.:Zusammenfassung I Abstract III Inhaltsverzeichnis V Abkürzungsverzeichnis VIII 1 Einleitung 1 1.1 Totholz als Bestandteil von Waldökosystemen 1 1.1.1 Vorkommen von Totholz 1 1.1.2 Klassifizierung von Totholz 1 1.1.3 Entstehung von Totholz 2 1.1.4 Totholz und Biodiversität 3 1.1.5 Totholz in Stoffkreisläufen 8 1.1.6 Totholz als wichtiges Strukturelement 9 1.2 Holzaufbau 10 1.2.1 Grundsätzlicher Aufbau von Holz 10 1.2.2 Der Lignozellulose-Komplex 14 1.3 Saprobionte Pilze als Spezialisten zur Überwindung der Ligninbarriere 18 1.3.1 Weißfäulepilze 18 1.3.2 Braunfäulepilze 20 1.3.3 Moderfäulepilze 22 1.4 Enzymatischer Ligninabbau 23 1.4.1 Laccase 23 1.4.2 Peroxidasen 26 1.5 Totholz - Stand der Forschung 33 1.5.1 Totholzabbau in Europa 33 1.5.2 Totholz und Waldbewirtschaftung 34 1.5.3 Abbauprozesse 34 1.5.4 Oxidative Enzyme im Totholz 36 2 Zielstellung der Arbeit 39 3 Methoden 43 3.1 Untersuchung von natürlichem Totholz auf den VIP-Flächen 43 3.1.1 Untersuchungsgebiet 43 3.1.2 Probenahme 47 3.1.3 Aufbereitung der Proben für die enzymatischen Messungen 49 3.1.4 Aktivitäten oxidativer Enzyme 50 3.1.5 Physikochemische Variablen der Totholzproben 52 3.1.6 Artenzusammensetzung der Pilze auf und im Totholz 54 3.1.7 Statistik 56 3.2 Erfassung der kleinräumigen Verteilung von Oxidoreduktasen in einem Totholzfragment 63 3.2.1 Probenahme 63 3.2.2 Untersuchung der Proben 65 3.2.3 Statistische Auswertung 66 3.3 Stickstoffexperiment 66 3.3.1 Experimentaufbau 66 3.3.2 Probenahme 68 3.3.3 Aufbereitung der Proben für die enzymatischen Messungen 69 3.3.4 Enzymatische Untersuchungen 69 3.3.5 Untersuchung mit markiertem Stickstoff 74 3.3.6 Statistische Analyse 74 3.4 Optimierung der organischen Extraktion in Vorbereitung der Ligninbestimmung 75 3.4.1 Methodisches Vorgehen 76 3.4.2 Ergebnisse zur Methodenentwicklung 78 3.4.3 Bewertung der Methodenentwicklung 80 4 Ergebnisse 83 4.1 Natürliches Totholz auf den VIP-Flächen 83 4.1.1 Totholzvariablen und Ihre Unterschiede zwischen den Baumarten 83 4.1.2 Einfluss der Waldbewirtschaftung auf die Variablen des Totholzabbaus 91 4.1.3 Veränderungen des Totholzes während der Zersetzung 92 4.1.4 Abhängigkeit der oxidativen Enzymaktivitäten von den physikochemischen Eigenschaften und den Pilzarten (OTUs) 99 4.1.5 Kleinräumige Verteilungsmuster der oxidativen Enzymaktivitäten in den Totholzstämmen 105 4.2 Kleinräumige Muster der oxidativen Enzymaktivitäten in einem einzelnen Totholzfragment 106 4.3 Stickstoffexperiment 111 5 Diskussion 115 5.1 Unterschiede im Zersetzungsprozess zwischen den Baumarten 115 5.2 Oxidative Enzymaktivitäten im Totholz 119 5.2.1 Bedeutung von Lacc, GenP und MnP für die Ligninmodifikation 119 5.2.2 Variabilität der Lacc-, GenP- und MnP-Aktivitäten 121 5.2.3 Kleinräumige Muster der Lacc-, GenP und MnP-Aktivitäten 122 5.2.4 Dynamik der oxidativen Enzymaktivitäten im Verlauf des Zersetzungsprozesses 123 5.2.5 Zusammenhänge zwischen den oxidativen Enzymaktivitäten und den Totholzvariablen 125 5.3 Veränderung des Totholzes über den Zersetzungsprozess 135 5.3.1 Die Artengemeinschaft 136 5.3.2 Die Holzbestandteile und der pH-Wert 138 5.3.3 Die Nährstoffe 139 5.4 Einfluss der Waldbewirtschaftung auf Variablen des Totholzabbaus 141 6 Ausblick 145 7 Thesen 151 8 Literaturverzeichnis 153 Anhang 169 A Charakteristik der Untersuchungsflächen 169 B NMDS-Ordination der pilzlichen Artengemeinschaft 172 C Daten der Totholzstämme 175 D Daten zu den Proben 177 E Daten zur Modellierung der Enzymaktivitäten und der Wahrscheinlichkeit, diese zu detektieren 178 F Daten zur Untersuchung des einzelnen F.-sylvatica-Totholzfragments 189 G Detailabbildungen zur Zersetzungsdynamik 192 H Semivariogrammdaten oxidativer Enzyme im Totholz der VIP-Flächen 195 I Km-Werte von Mangan-Peroxidasen (MnP) für Mangan(II)-Ionen (Mn2+) aus der Literatur 196 J Zuordnung der Fäuletypen zu den Pilzarten 198 K Publikationen 208 L Danksagung 251 M Rechtliche Erklärung 253 In forest ecosystems, deadwood is an important component that provides habitat and substrate for numerous organisms, contributes to the carbon and nutrient cycle as well as serves as a structural element. Overcoming the lignin barrier is a key process in deadwood degradation. Only specialized saprotrophic fungi of the phyla Basidiomycota and Ascomycota developed different strategies – the rot types – to degrade lignin or to modify it in way, which allows them to get access to the polysaccharides (cellulose and hemicelluloses) that are incrusted within the lignocellulosic complex. In this context, basidiomycetous white rot fungi secreting oxidative enzymes (especially laccases and peroxidases) are of particular importance, since they are the only organisms that are able to substantially mineralize lignin to carbon dioxide (CO2). Although lignin degradation is such an important process for deadwood degradation, oxidative enzyme activities have been only poorly studied under n...
    Keywords: Info:Eu-Repo/Classification/Ddc/570 ; Ddc:570 ; Lignin; Pilze; Enzym; Nährstoff; Forstwirtschaft; Holz; Laccase; Mangan-Peroxidase; Peroxidasen; Holzfäule; Wasserstoffionenkonzentration; Fichte; Waldkiefer; Rotbuche ; Saprobionte Pilze ; Oxidative Enzyme ; Nährstoffe ; Lignin ; Waldbewirtschaftungsintensität ; Starkes Totholz ; Laccase ; Mangan-Peroxidase ; Generelle Peroxidase ; Holzfäule ; Weißfäule ; Braunfäule ; Moderfäule ; Ph-Wert ; Holz ; Buche ; Fichte ; Kiefer ; Extraktive ; Pilzliche Artengemeinschaft ; Saproxylic Fungi ; Oxidative Enzyme ; Nutrients ; Lignin ; Forest Management Intensity ; Coarse Woody Debris ; Laccase ; Manganese Peroxidase ; General Peroxidase ; Wood Rot ; White Rot ; Brown Rot ; Soft Rot ; Ph ; Wood ; Beech ; Spruce ; Pine ; Extractives ; Fungal Community
    Source: Networked Digital Library of Theses and Dissertations
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