Carbon and Nitrogen Cycling in European Forest Ecosystems

520 Pages·2000·37.462 MB·English

by  E.-D. Schulze (auth.), Prof. Dr. Ernst-Detlef Schulze (eds.)

Ecological Studies, Vol. 142
Analysis and Synthesis
Edited by
M.M. Caldwell, Logan, USA
G. Heldmaier, Marburg, Germany
0.1. Lange, Wiirzburg, Germany
H.A. Mooney, Stanford, USA
E.-D. Schulze, Jena, Germany
U. Sommer, Kiel, Germany
Ecological Studies
Volumes published since 1994 are listed at the end of this book.
Springer-Verlag Berlin Heidelberg GmbH
E.-D. Schulze (Ed.)
Carbon and Nitrogen
Cycling in European
Forest Ecosystems
With 183 Figures, 14 in Color, 106 Tables, and CD-ROM
Springer
Prof. Dr. Ernst-Detlef Schulze
Max-Planck-Institute for Biogeochemistry
P.O. Box 10 01 64
07701 Jena
Germany
Design of the cover illustration by Barbara Lühker
Additional material to this book can be downloaded from http://extras.springer.com.
ISSN 0070-8356
ISBN 978-3-540-67239-5
Library of Congress Cataloging-in-Publication Data
Carbon and nitrogen cycling in European forest ecosystems /E.-D. Schulze (ed.).
p. cm.-(Ecological studies, ISSN 0070-8356; vol. 142)
Includes bibliographical references.
ISBN 978-3-540-67239-5 ISBN 978-3-642-57219-7 (eBook)
DOI 10.1007/978-3-642-57219-7
I. Carbon cycle (Biogeochemistry) -Europe. 2. Nitrogen cycle - Europe. 3. Forest
ecology-Europe. I. Schulze, E.-D. (Ernst-Detlef), 1941. II. Ecological studies; v. 142.
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Preface
One might think that the principles of carbon and nitrogen cycles of forests are text
book knowledge. However, if we are asked to quantify the fluxes and to make firm
estimates about relations to natural and human-induced disturbances, then our
answers will rapidly become very vague. This ecological dilemma becomes even more
obvious, when scientists are questioned by politicians to verifiably state whether a
region is a source or a sink for carbon. The Kyoto protocol of the Climate Conven
tion poses this question to the scientific community, and the answers are contradic
tory. Fan et al. [Science 282:442(1998)] claimed that the Northern Hemisphere
sink for carbon is located solely along the continental US, while Ciais 0. Geophys.
Res., 1999) located the Northern Hemisphere sink in Siberia, and Lloyd (Functional
Ecology 1999) convincingly showed that the main sink is located in the tropics.
Thus, we are almost lost and, as soon as we depart from natural vegetation into
managed systems, the uncertainties are even greater.
Based on a 5-year investigation, this books, adopts the IGBP approach of studying
continental transects, because only by comparison beyond our "backyard" we are able
to identify general principles. The study of transects seems to be the most manage
able approach to quantify ecosystem processes on a continental scale, which is needed
if ecology wants to compete and co-operate with earth system science in the future.
Thus, the demanding task was undertaken to study whole ecosystems in a compara
tive way, and under a joint protocol along a transect reaching from north Sweden to
central Italy. It is quite clear that it remains impossible to study a geographic transect
in a contiguous way. In the present work, the transect was used as a tool in order to
maintain certain conditions as constant as possible, e.g. soils or type of vegetation.
The focus was rather on other important variables (e.g. climate and deposition), and
to determine processes that drive the ecosystem mass balance of carbon and nitro
gen along these conditions. This task became further complicated by the insight (see
Schulze and Mooney, Ecological Studies 99) that the biogeochemical cycles of terres
trial ecosystems can no longer be studied without considering the specific function
of biodiversity. Obviously, European forests are not very diverse with respect to tree
species. In fact, they are managed mono cultures, but a wealth of species remains in
the soil, and the subtle interaction between plants and mycorrhizae, and other soil
organisms is not at all obvious. This book addresses this problem and investigates to
what extent this diversity affects ecosystem functioning, and why.
In view of the present request by the public and to obtain answers that were posed
by the Kyoto protocol, we would like to acknowledge the foreseeing actions of the EU
to emphasise transect studies across Europe, and to foster basic understanding of
processes. When the project started, the aim to support basic research was a conse
quence of the problems experienced with identifying the causes of forest decline. In
VI Preface
view of the present demand for data on carbon sequestration, the early decision by
the EU may demonstrate the importance of basic science for the future of a society.
We would like to make this point at a time when scientists are increasingly asked to
carry out research of public concern, or with direct applications.
Jena, June 2000 E.-D. Schulze
Contents
Part A Introduction to the European Transect
1 The Carbon and Nitrogen Cycle of Forest Ecosystems
E.-D. Schulze .............................................. 3
1.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2 The Carbon and Nitrogen Cycles .............................. 3
1.3 The NIPHYS/CANIF Project .................................. 8
1.4 Experimental Design ....................................... 10
1.5 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11
References ...................................................... 11
2 Experimental Sites in the NIPHYS/CANIF Project
T. Persson, H. van Oene, A.F. Harrison, P.S. Karlsson, G.A. Bauer,
J. Cerny, M.-M. Couteaux, E. Dambrine, P. Hogberg, A. Kj0ller,
G. Matteucci, A. Rudebeck, E.-D. Schulze, and T. Paces. . . . . . . . . . . . . .. 14
2.1 Site Description. The NIPHYS/CANIF Transect ................... 14
2.2 Soil Characteristics ......................................... 26
2.3 Ecosystem C and N Pools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 36
2.4 Database. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 39
2.5 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 42
References ...................................................... 46
Part B Plant-Related Processes
3 Tree Biomass, Growth and Nutrient Pools
G. Scarascia-Mugnozza, G.A. Bauer, H. Persson, G. Matteucci,
and A. Masci .............................................. 49
3.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49
3.2 Experimental Background .................................... 50
3.3 Biomass. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52
3.4 Forest Productivity ......................................... 54
3.5 Carbon and Nutrient Pools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56
3.6 Allometric and Functional Relations ........................... 58
3.7 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
References ..................................................... 62
VIII Contents
4 Linking Plant Nutrition and Ecosystem Processes
G.A. Bauer, H. Persson, T. Persson, M. Mund, M. Hein,
E. Kummetz, G. Matteucci, H. van Oene, G. Scarascia-Mugnozza,
and E.-D. Schulze .......................................... 63
4.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
4.2 Experimental Approach .................................... 64
4.3 Nutrient Concentrations .................................... 65
4.4 Nutrient Contents ......................................... 73
4.5 Nitrogen Partitioning in Different Tree Compartments ........... 82
4.6 Ecosystem C and N Pools ... ................................ 88
4.7 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
References ..................................................... 95
5 Root Growth and Response to Nitrogen
C. Stober, E. George, and H. Persson . . . . . . . . . . . . . . . . . . . . . . . . . . 99
5.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
5.2 Approaches to the Study of Root Growth ....................... 100
5.3 Root Growth Measurements Obtained by Soil Coring ............. 105
5.4 Root Growth Measurements Obtained by Root Windows .......... 107
5.5 Root Growth Measurements Obtained by In-Growth Cores ......... 113
5.6 Root Growth at Different European Forest Sites ................. 116
5.7 Conclusions.. . .. . . . . .. . . . . .. . . .. . .. . .. . . . . . .. . .. . . . .. . . .. 118
References ..................................................... 119
6 Nitrogen Uptake Processes in Roots and Mycorrhizas
T. Wallenda, C. Stober, L. Hogbom, H. Schinkel, E. George,
P. Hogberg, and D.J. Read ................................... 122
6.1 Introduction ............................................. , 122
6.2 Approaches to Study Different Aspects of the N Uptake Process ..... 123
6.3 Studies with Excised Roots and Mycorrhizas .................... 125
6.4 Field-Based Experiments ................................... 135
6.5 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 139
References ..................................................... 141
7 The Fate of 15N-Labelled Nitrogen Inputs to Coniferous and
Broadleaf Forests
G. Gebauer, B. Zeller, G. Schmidt, C. May, N. Buchmann, M. Colin-
Belgrand, E. Dambrine, F. Martin, E.-D. Schulze, and P. Bottner ...... 144
7.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 144
7.2 Sites of Investigation ....................................... 145
7.3 Approaches to Study the Fate of lsN-Labelled Nitrogen Inputs ...... 147
7.4 N Release and Tree Uptake from lsN-Labelled Decomposing
Litter in a Beech Forest in Aubure ............................. ISO
7.5 Ecosystem Partitioning of lsN-Labelled Ammonium
and Nitrate on the Sites in the Fichtelgebirge and Steigerwald ...... 157
7.6 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 166
References ..................................................... 168
Contents IX
8 Canopy Uptake and Utilization of Atmospheric Pollutant Nitrogen
A.F. Harrison, E.-D. Schulze, G. Gebauer, and G. Bruckner .......... 171
8.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 171
8.2 Atmospheric Nitrogen Pollutants ............................. 172
8.3 Pathways for Canopy Uptake of Nitrogen ...... . . . . . . . . . . . . . . .. 174
804 Approaches to the Determination of Canopy Uptake of Nitrogen .... 175
8.5 Review of Research ........................................ 177
8.6 Role in the Critical Load .................................... 182
8.7 Ecophysiological Consequences of Canopy N Uptake ............. 182
8.8 Conclusions ............................................ , 183
8.9 Way Forward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 183
8.10 Policy Implications ........................................ 184
References ..................................................... 184
9 Biotic and Abiotic Controls Over Ecosystem Cycling
of Stable Natural Nitrogen, Carbon and Sulphur Isotopes
G.A. Bauer, G. Gebauer, A.F. Harrison, P. Hogberg, L. Hogbom,
H. Schinkel, A.F.S. Taylor, M. Novak, F. Buzek, D. Harkness, T. Persson,
and E.-D. Schulze ......................................... 189
9.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 189
9.2 Approaches to the Study of Stable Isotopes in the Field ........... 189
9.3 81sN of Ammonium and Nitrate in Wet Deposition ............... 191
904 Stable Isotope Signatures in Different Ecosystem Compartments .... 197
9.5 815N Signatures as Indicators
of N Saturation in Forest Ecosystems .......................... 207
9.6 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 211
References ..................................................... 213
Part C Heterotrophic Processes
10 Soil Respiration in Beech and Spruce Forests in Europe:
Trends, Controlling Factors, Annual Budgets and Implications
for the Ecosystem Carbon Balance
G. Matteucci, S. Dore, S. Stivanello, C. Rebmann, and N. Buchmann. .. 217
10.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 217
10.2 Approaches to Measuring Soil Respiration ...................... 218
10.3 Daily and Seasonal Trends in Soil Respiration
and Climatic Variables ..................................... 221
IDA Factors Controlling Soil Respiration ........................... 226
10.5 Comparison of Chamber Measurements
with the Eddy Covariance Measurements Below the Canopy ........ 230
10.6 Annual Budgets of Soil Respiration ........................... 231
10.7 Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 233
References ..................................................... 234