[Advances in Ecological Research] Litter Decomposition: A Guide to Carbon and Nutrient Turnover Volume 38 || Contents

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ContentsPreface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vAcknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viiIntroductionI. General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1A. Decomposition, Nutrient Turnover, and GlobalClimate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3B. Biomass Distribution between Soil and Above-GroundEcosystem Compartments . . . . . . . . . . . . . . . . . . . . . . . 9C. The Importance of Balance . . . . . . . . . . . . . . . . . . . . . . 12Litter FallI. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20II. Litter Fall AmountsMain Patterns and Regulating Factors . 21A. Patterns on the Forest Stand Level . . . . . . . . . . . . . . . . 21B. Litter Fall Patterns in Scots PineA Case Study . . . . . . 23III. A Model for Accumulated Litter Fall, Stand Level . . . . . . . . 26A. General Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26B. A Case Study for a Scots Pine Stand . . . . . . . . . . . . . . . 26IV. Main Litter-Fall Patterns on a Regional Level: Scots Pine andNorway Spruce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28A. Distribution of Species . . . . . . . . . . . . . . . . . . . . . . . . . 28B. Factors Influencing Amounts of Litter Fall. . . . . . . . . . . 28C. Needle Litter FallPattern and Quantities: Scots Pineand Other Pine Species . . . . . . . . . . . . . . . . . . . . . . . . . 29D. Basal Area and Canopy Cover. . . . . . . . . . . . . . . . . . . . 35E. Needle Litter Quantities: Norway Spruce . . . . . . . . . . . . 36F. Comparison of and Combination of Species . . . . . . . . . . 36G. Litter Fall on a Continental to Semiglobal Scale . . . . . . . 37V. The Fiber Structure and OrganicChemical Components ofPlant Litter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40A. The Fiber. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40B. The OrganicChemical Components. . . . . . . . . . . . . . . . 43x CONTENTSVI. Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46A. General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46B. The Trees Withdraw Nutrients before Shedding theirFoliar Litter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49C. Scots PineA Case Study. . . . . . . . . . . . . . . . . . . . . . . 53D. Foliar Litter N Concentration in a Trans-EuropeanTransect, Several Species. . . . . . . . . . . . . . . . . . . . . . . . 58E. Several Deciduous and Coniferous Leaf Litters. . . . . . . . 58VII. Anthropogenic Influences . . . . . . . . . . . . . . . . . . . . . . . . . . 62A. Nitrogen-Fertilized Scots Pine and NorwaySpruce Monocultures . . . . . . . . . . . . . . . . . . . . . . . . . . 62B. The EVect of Heavy Metal Pollution . . . . . . . . . . . . . . . 67VIII. Methods for Litter Collection . . . . . . . . . . . . . . . . . . . . . . . 69A. Quantities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69B. Qualitative Sampling . . . . . . . . . . . . . . . . . . . . . . . . . . 71Decomposers: Soil Microorganisms and AnimalsI. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73II. Communities of Soil Microorganisms and Animals . . . . . . . . 75A. Soil Microorganisms. . . . . . . . . . . . . . . . . . . . . . . . . . . 75B. Soil Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77III. The Degradation of the Main Polymers in Plant Fibers . . . . . 79A. Degradation of Cellulose . . . . . . . . . . . . . . . . . . . . . . . 79B. Degradation of Hemicelluloses . . . . . . . . . . . . . . . . . . . 82C. EVects of N, Mn, and C Sources on the Degradationof Lignin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83D. Degradation of Lignin . . . . . . . . . . . . . . . . . . . . . . . . . 87IV. Degradation of Fibers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92A. Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92B. Bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93V. Microbial Communities and the Influence of Soil Animals. . . 94A. Microbial Succession and Competition. . . . . . . . . . . . . . 94B. EVects of Soil Animals on the Decomposition Process . . 96Changes in Substrate Composition and Rate-RegulatingFactors during DecompositionI. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . 102II. OrganicChemical Changes During Litter Decomposition . . . 104A. Decomposition of Single Chemical Components andGroups of Compounds . . . . . . . . . . . . . . . . . . . . . . . . . 104B. Relationships between Holocellulose and Ligninduring Decomposition . . . . . . . . . . . . . . . . . . . . . . . . . 114CONTENTS xiIII. Concentrations of Nutrients and Heavy Metals DuringLitter Decay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114A. Nitrogen (N) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116B. Phosphorus (P) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116C. Sulphur (S). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116D. Potassium (K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117E. Calcium (Ca) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117F. Magnesium (Mg) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117G. Other Metals and Heavy Metals inNatural Concentrations . . . . . . . . . . . . . . . . . . . . . . . . . 118IV. A Three-Phase Model Applied to Litter of DiVerent InitialChemical Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119A. Overview of the Model . . . . . . . . . . . . . . . . . . . . . . . . . 119B. Initial Decomposition Rates for Newly Shed LitterTheEarly Decomposition Stage . . . . . . . . . . . . . . . . . . . . . . 119C. Decomposition in the Late StageA Phase Regulatedby Lignin Decomposition . . . . . . . . . . . . . . . . . . . . . . . 129D. Link between the Retardation of Litter Decomposition,Lignin Degradation Rate and N Concentration. . . . . . . . 137E. Comments on Spruce Needle Litter Decompositionversus the Three-Phase Model . . . . . . . . . . . . . . . . . . . . 139F. The Litter Close to the Limit Value and at aHumus-Near Stage . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142G. Do Limit Values Indicate a Stop in the LitterDecomposition Process? . . . . . . . . . . . . . . . . . . . . . . . . 150V. Lignin Dynamics in Decomposing Litter. . . . . . . . . . . . . . . . 150A. Repeatability of Patterns in LigninConcentration Changes . . . . . . . . . . . . . . . . . . . . . . . . . 150B. Variation in the Increase in Lignin ConcentrationRelative to DiVerent Initial Lignin Concentrations inthe Litter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153C. Variation in Lignin Concentration Increase Rateas Compared to DiVerent Concentrationsof N in Litter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153VI. Does the Litter Chemical Composition Influence Leachingof Compounds from Decomposing Litter?. . . . . . . . . . . . . . . 154xii CONTENTSNitrogen Dynamics in Decomposing LitterI. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157II. The Dynamics of NitrogenThree Phases inDecomposing Litter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159A. General Comments. . . . . . . . . . . . . . . . . . . . . . . . . . . . 159B. The Leaching Phase . . . . . . . . . . . . . . . . . . . . . . . . . . . 161C. Nitrogen Accumulation PhaseA Phase with a NetUptake and a Retention of N . . . . . . . . . . . . . . . . . . . . 164D. A Release Mechanism. . . . . . . . . . . . . . . . . . . . . . . . . . 170E. The Final Release Phase . . . . . . . . . . . . . . . . . . . . . . . . 176III. Nitrogen Concentration Versus AccumulatedLitter Mass Loss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177IV. Nitrogen Concentration in Litter Decomposing to theLimit Value and in Humus . . . . . . . . . . . . . . . . . . . . . . . . . 181A. Background and Some Relationships . . . . . . . . . . . . . . . 181B. A Model and a Case Study for Calculating NConcentrations in Humus . . . . . . . . . . . . . . . . . . . . . . . 182Origin and Structure of Secondary Organic Matter andSequestration of C and NI. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . 185II. Terminology According to Traditional Humus Classificationand Chemical Composition of Secondary Organic Matter . . . 189III. Origin of Secondary Organic MatterSomePrimary Scenarios. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194A. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . 194B. Two Traditional Scenarios . . . . . . . . . . . . . . . . . . . . . . 195C. Some More Recent Approaches to Humic Substances . . . 196IV. The Role of SOM in Soil . . . . . . . . . . . . . . . . . . . . . . . . . . 198V. What Litter Components May Be of Importance for theFormation of Humus?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200VI. The Accumulation Rate of Humus. . . . . . . . . . . . . . . . . . . . 203A. Direct Measurements of Humus Accumulation. . . . . . . . 203B. Accumulation of HumusEstimates . . . . . . . . . . . . . . . 204C. How Reliable are Quantitative Estimates ofHumus Accumulation? . . . . . . . . . . . . . . . . . . . . . . . . . 210VII. May All Humus be Decomposed or Just a Fraction?. . . . . . . 210A. DiVerent FractionsGeneral Comments . . . . . . . . . . . . 210B. Four Cases of Turnover of Humus Layers . . . . . . . . . . . 211CONTENTS xiiiVIII. Humus Accumulation and Decomposition Versus TheConcept Steady State . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215A. Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215B. Why Is It an Error to Use the Concept Steady State? . 216IX. Nitrogen Sequestration to SOM . . . . . . . . . . . . . . . . . . . . . . 217A. We Can Estimate the Sequestration Rate of N inStable Organic Matter. . . . . . . . . . . . . . . . . . . . . . . . . . 217B. We Can Validate the Long-Term Accumulation ofStable Nitrogen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218X. The Capacity of SOM to Store N. . . . . . . . . . . . . . . . . . . . . 221XI. Can DiVerent Capacities to Sequester N Be Related toSpecies or to The Initial Litter N Concentration? . . . . . . . . . 222XII. How Stable Is the Long-term N Stored in Humus? . . . . . . . . 225Climatic and Geographic Patterns in DecompositionI. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227II. The Microbial Response to Temperature and Moisture . . . . . 228III. The Influence of Climate on Early-Stage Decompositionof Scots Pine Needle Litter . . . . . . . . . . . . . . . . . . . . . . . . . 229A. Early-Stage Decomposition at One ForestStand over Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229B. Decomposition Studies in Transects with ScotsPine and Norway Spruce . . . . . . . . . . . . . . . . . . . . . . . . 231IV. The EVect of Substrate Quality on Mass-Loss Ratesin Scots Pine Transects . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240A. Early Stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240B. Decomposition over a Transect with Scots PineMonoculturesThe Late Stage . . . . . . . . . . . . . . . . . . . 242C. Respiration from Humus from Scots PineStands in a Pan-European Transect . . . . . . . . . . . . . . . . 245V. The Influence of Climate on Decomposition of NorwaySpruce Litter in a Transect . . . . . . . . . . . . . . . . . . . . . . . . . 250A. General Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250B. Climate Versus First-Year Mass Loss . . . . . . . . . . . . . . . 251C. Lignin-Mediated EVects on Litter DecompositionRates during Late Stages of Decomposition . . . . . . . . . . 252VI. A Series of Limiting Factors for Decomposing Litter. . . . . . . 255A. Factors Influencing Lignin Degradation Rates . . . . . . . . 255VII. The Influence of Climate on Decomposition of Root Litter . . 257xiv CONTENTSVIII. Litter Chemical Changes as Related to Climate. . . . . . . . . . . 259A. Development of Litter N Concentration withClimate in Decomposing Scots Pine NeedleLitter (Transects I and II) . . . . . . . . . . . . . . . . . . . . . . . 259B. Development of Litter Lignin Concentration withClimate in Decomposing Needle Litter. . . . . . . . . . . . . . 260Anthropogenic Impacts on Litter Decomposition and SoilOrganic MatterI. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . 263II. Fate of Pollutants in Litter and Soil. . . . . . . . . . . . . . . . . . . 264A. General Background. . . . . . . . . . . . . . . . . . . . . . . . . . . 264B. Acidic Precipitation . . . . . . . . . . . . . . . . . . . . . . . . . . . 265C. Heavy Metals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266D. Accumulation of Heavy Metals in DecomposingLitterA Case Study . . . . . . . . . . . . . . . . . . . . . . . . . . 268E. Sources of Heavy Metals in Litter . . . . . . . . . . . . . . . . . 271F. Organic Pollutants . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275III. EVects of Pollutants on Decomposition . . . . . . . . . . . . . . . . 277A. Heavy Metals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277B. Acidic Precipitation . . . . . . . . . . . . . . . . . . . . . . . . . . . 280C. Organic Pollutants . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281D. EVects of Climate Change. . . . . . . . . . . . . . . . . . . . . . . 283E. Changes in Water Regimen . . . . . . . . . . . . . . . . . . . . . . 289Methods in Studies of Organic Matter DecayI. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . 291II. Incubation Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292A. In Situ (Field) Methods . . . . . . . . . . . . . . . . . . . . . . . . 292B. Decomposition RateLaboratory Methods . . . . . . . . . . 309III. Studying Chemical Changes During Decomposition . . . . . . . 314A. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . 314B. Preparation of Samples for Chemical Analysisand Some Analytical Techniques . . . . . . . . . . . . . . . . . . 315IV. Data Analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320A. Regression Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . 320B. Analysis of Variance (ANOVA) . . . . . . . . . . . . . . . . . . 324C. Multivariate Methods . . . . . . . . . . . . . . . . . . . . . . . . . . 326V. Presentation of the Results . . . . . . . . . . . . . . . . . . . . . . . . . 328CONTENTS xvAppendix I. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333Appendix II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407Cumulative List of Titles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423