Litter decomposition a guide to carbon and nutrient turnover (advances in ecological research, volume 38)

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The present book is written mostly with students and teachers in mind. The authors hope that this book will be useful at all levels of study, from general ecology courses, where decomposition processes often are covered briefly, through more advanced courses in ecosystems ecology, soil ecology, and biogeochemistry, where at least some deeper aspects of organic matter decay should be covered, ending with courses for graduate students who decide to take the first step in their research careers in this topic.

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  • Advances in ECOLOGICAL RESEARCH VOLUME 38
  • Advances in Ecological Research Series Editor: HAL CASWELL Biology Department Woods Hole Oceanographic Institution Woods Hole, Massachusetts
  • Advances in ECOLOGICAL RESEARCH VOLUME 38 LITTER 2005 BJRN BERG AND RYSZARD LASKOWSKI DECOMPOSITION: A GUIDE TO CARBON AND NUTRIENT TURNOVER
  • Elsevier Academic Press 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobalds Road, London WC1X 8RR, UK This book is printed on acid-free paper. Copyright 2005, Elsevier Ltd. except: Chapter 1, Food Webs, Body Size, and Species Abundance in Ecological Community Description, copyright 2005 by Tomas Jonsson, Joel E. Cohen, and Stephen R. Carpenter. Chapter 3, Estimating Relative Energy Fluxes Using the Food Web, Species Abundance, and Body Size, copyright 2005 by Daniel C. Reuman and Joel E. Cohen. Elsevier reserves all rights to the portions copyrighted by Elsevier. The authors reserve all rights to the copyrighted by them. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the Publisher. The appearance of the code at the bottom of the first page of a chapter in this book indicates the Publishers consent that copies of the chapter may be made for personal or internal use of specific clients. This consent is given on the condition, however, that the copier pay the stated per copy fee through the Copyright Clearance Center, Inc. (www.copyright.com), for copying beyond that permitted by Sections 107 or 108 of the U.S. Copyright Law. This consent does not extend to other kinds of copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collective works, or for resale. Copy fees for pre-2005 chapters are as shown on the title pages. If no fee code appears on the title page, the copy fee is the same as for current chapters. 0065-2504/2005 $35.00 Permissions may be sought directly from Elseviers Science & Technology Right Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: permissions@elsevier.com.uk. You may also complete your request on-line via the Elsevier homepage (http://elsevier.com), by selecting Customer Support and then Obtaining Permissions. For all information on all Elsevier Academic Press publications visit our Web site at www.books.elsevier.com ISBN: 0-12-013938-3 PRINTED IN THE UNITED STATES OF AMERICA 0 5 06 07 08 9 8 7 6 5 4 3 2 1
  • Preface The idea of this book, which we are able to oVer you now, thanks to Elsevier, was born several years ago when we were working together on review articles summarizing knowledge on litter decomposition, nutrient dynamics, and humus buildup in forests of the Northern Hemisphere. After years of study- ing organic matter decomposition, we considered ourselves to have good insight into the progress in this branch of ecology/biogeochemistry, and it was not diYcult to notice that, although intensively studied by a number of research teams all over the world, the subject was not very well represented on the bookshelves. Virtually no single comprehensive book devoted to this subject had been published for a long time, and, in fact, very few have ever been published (see the References at the end of this volume). To both of usteaching ecology as well as more specialized courses in soil ecology and ecotoxicologythis situation was not merely unsatisfactory, considering the importance of decomposition processes for almost every aspect of life on earth, but also very inconvenient for our students who did not have any source summarizing the state of current research in the discipline. When you lack a proper handbook, you must write one yourself and we decided to do just that. Although many years have passed from the birth of the idea until we could submit the manuscript, not much has changed in the general market. One notable exception is the book coauthored by one of us and published by SpringerVerlag in 2003. However, while that book is directed toward specialists, the present one has been written mostly with students and teachers in mind. We hope that this book will be useful at all levels of study, from general ecology courses, where decomposition process- es often are covered briey, through more advanced courses in ecosystems ecology, soil ecology, and biogeochemistry, where at least some deeper aspects of organic matter decay should be covered, ending with courses for graduate students who decide to take the rst step in their research careers in this topic. While teachers and students in more general subjects will nd the most basic information on decomposition processes in this book, we hope that scientists and graduate students working on decomposition processes will be satised with the more detailed information and the overview of the latest publications on the topic as well as the methodological chapter where
  • practical information on methods useful in decomposition studies can be found. We hope that university teachers like us will nd the book useful in preparing their courses. In particular, those who do not specialize in decom- position studies should nd a wealth of knowledge gathered in one, relatively compact volume. A useful addition for classes and selfteaching is Appendix II, with real research data and an Internet link that can be used for learning diVerent statistical techniques mentioned in the book or even for organizing minor research projects without the necessity of spending long years on eld studies, which, in most cases, is simply impossible during regular courses. Of course, we do not believe that our book will satisfy the needs of everyone. Throughout the book, we have had to nd a balance between completeness of the knowledge presented and compactness of particular chapters. We realize that our personal opinion on what is the best tradeoV was not necessarily optimal in all cases. Therefore, we will be happy to hear your opinions and suggestions. If the book appears useful, there is the possibility of publishing an updated version in a few years. Our email addresses are given below this Preface: you may be certain that every message will be carefully read and thought through. E-mail address: Bjorn Berg: BBE@kvl.dk, 0708212424@euromail.se Ryszard Laskowski: r.laskowski@eko.uj.edu.pl vi PREFACE
  • Acknowledgements This book would never have been written without generations of students who participated actively in our courses in general ecology, soil ecology, and ecotoxicology at the Jagiellonian University in Krakow, Poland. Further, graduate students and colleagues at the University of Bayreuth, Germany, and Oregon State University were helpful at diVerent stages of writing the book. Ryszard Laskowski: A number of our colleagues should be acknowledged because, without their encouragement and help, we would never have under- taken the challenge of writing this handbook. Professor Wadysaw Grodzinski, the late head of the Department of Ecosystem Studies at the Jagiellonian University, was the rst who turned my research interests toward litter decomposition studies and led the rst research projects on this subject at the Jagiellonian University. We had the great pleasure to work together in a number of projects with Professor Krystyna Grodzinska, Head of the Department of Botany, Polish Academy of Sciences. Her knowledge matches her personal charm and friendliness, and it is hard for me to imagine my scientic career without her help and cooperation. Among those without whom this book would probably never have come into being is January Weiner, professor and Head of the Department of Ecosys- tem Studies at the Jagiellonian University. No other person has ever oVered me so much encouragement and taught me so much about science in general. Finally, I express my greatest gratitude to my colleagues from the Depart- ment of Ecotoxicology, Jagiellonian University: Paulina Kramarz, Maciej Maryanski, and Maria Niklinska, who helped me in my research for many years. Particularly Maria and Maciej spent countless hours on our common research on litter decomposition in European forests. The joy of common eldwork, long days and nights spent on chemical analyses, the excitement of new ndings is unforgettable. Bjorn Berg: I want to thank Professor C. O. Tamm for all his support of my work, both within the SWECON project and after, allowing a period of no fewer than 18 years to be devoted to work on litter decomposition. During the same period, I had really skilled, not to say fantastic, assistance from my three laboratory assistants, Annette Ewertsson, Birgitta Holm, and AnnSo Pettersson. The patient preparation and cleaning work of
  • hundreds of thousands of litterbags at Jadraas Experimental Park resulted in the creation of a large database that allowed, among other work, this book to be written. The support of Senior Scientist Per Gundersen during the European Union CNTER project (Contract number QLK5200100596) and during my stay as a guest scientist at the Center for Forest Landscape and Planning, KVL, Horsholm, Denmark, was essential for this book to be written. I also thank Professor Egbert Matzner of the institute BITO K, University of Bayreuth, Germany, for his support of this book. Copenhagen, August 2005 Bjorn Berg Krakow, August 2005 Ryszard Laskowski viii ACKNOWLEDGEMENTS
  • Contents Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Introduction I. General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 A. Decomposition, Nutrient Turnover, and Global Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 B. Biomass Distribution between Soil and Above-Ground Ecosystem Compartments . . . . . . . . . . . . . . . . . . . . . . . 9 C. The Importance of Balance . . . . . . . . . . . . . . . . . . . . . . 12 Litter Fall I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 II. Litter Fall AmountsMain Patterns and Regulating Factors . 21 A. Patterns on the Forest Stand Level . . . . . . . . . . . . . . . . 21 B. Litter Fall Patterns in Scots PineA Case Study . . . . . . 23 III. A Model for Accumulated Litter Fall, Stand Level . . . . . . . . 26 A. General Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 B. A Case Study for a Scots Pine Stand . . . . . . . . . . . . . . . 26 IV. Main Litter-Fall Patterns on a Regional Level: Scots Pine and Norway Spruce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 A. Distribution of Species . . . . . . . . . . . . . . . . . . . . . . . . . 28 B. Factors Inuencing Amounts of Litter Fall. . . . . . . . . . . 28 C. Needle Litter FallPattern and Quantities: Scots Pine and Other Pine Species . . . . . . . . . . . . . . . . . . . . . . . . . 29 D. Basal Area and Canopy Cover. . . . . . . . . . . . . . . . . . . . 35 E. Needle Litter Quantities: Norway Spruce . . . . . . . . . . . . 36 F. Comparison of and Combination of Species . . . . . . . . . . 36 G. Litter Fall on a Continental to Semiglobal Scale . . . . . . . 37 V. The Fiber Structure and OrganicChemical Components of Plant Litter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 A. The Fiber. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 B. The OrganicChemical Components. . . . . . . . . . . . . . . . 43
  • VI. Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 A. General Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 B. The Trees Withdraw Nutrients before Shedding their Foliar Litter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 C. Scots PineA Case Study. . . . . . . . . . . . . . . . . . . . . . . 53 D. Foliar Litter N Concentration in a Trans-European Transect, Several Species. . . . . . . . . . . . . . . . . . . . . . . . 58 E. Several Deciduous and Coniferous Leaf Litters. . . . . . . . 58 VII. Anthropogenic Inuences . . . . . . . . . . . . . . . . . . . . . . . . . . 62 A. Nitrogen-Fertilized Scots Pine and Norway Spruce Monocultures . . . . . . . . . . . . . . . . . . . . . . . . . . 62 B. The EVect of Heavy Metal Pollution . . . . . . . . . . . . . . . 67 VIII. Methods for Litter Collection . . . . . . . . . . . . . . . . . . . . . . . 69 A. Quantities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 B. Qualitative Sampling . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Decomposers: Soil Microorganisms and Animals I. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 II. Communities of Soil Microorganisms and Animals . . . . . . . . 75 A. Soil Microorganisms. . . . . . . . . . . . . . . . . . . . . . . . . . . 75 B. Soil Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 III. The Degradation of the Main Polymers in Plant Fibers. . . . . 79 A. Degradation of Cellulose . . . . . . . . . . . . . . . . . . . . . . . 79 B. Degradation of Hemicelluloses . . . . . . . . . . . . . . . . . . . 82 C. EVects of N, Mn, and C Sources on the Degradation of Lignin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 D. Degradation of Lignin . . . . . . . . . . . . . . . . . . . . . . . . . 87 IV. Degradation of Fibers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 A. Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 B. Bacteria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 V. Microbial Communities and the Inuence of Soil Animals. . . 94 A. Microbial Succession and Competition. . . . . . . . . . . . . . 94 B. EVects of Soil Animals on the Decomposition Process . . 96 Changes in Substrate Composition and Rate-Regulating Factors during Decomposition I. Introductory Comments . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 II. OrganicChemical Changes During Litter Decomposition . . . 104 A. Decomposition of Single Chemical Components and Groups of Compounds . . . . . . . . . . . . . . . . . . . . . . . . . 104 B. Relationships between Holocellulose and...

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