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Soil Respiration and the Environment
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Soil Respiration and
the Environment
Yiqi Luo and Xuhui Zhou
AMSTERDAM • BOSTON • HEIDELBERG

LONDON • NEW YORK • OXFORD
PARIS • SAN DIEGO • SAN FRANCISCO
SINGAPORE • SYDNEY • TOKYO
Academic Press is an imprint of Elsevier
Academic Press is an imprint of Elsevier
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525 B Street, Suite 1900, San Diego, California 92101-4495, USA
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Copyright © 2006, Elsevier, Inc. All rights reserved.
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.
Permissions may be sought directly from Elsevier’s Science & Technology Rights Department
in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: permissions@
elsevier.com. You may also complete your request on-line via the Elsevier homepage (http://
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“Obtaining Permissions.”
Library of Congress Cataloging-in-Publication Data
British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library.
ISBN 13: 978-0-12-088782-8

ISBN 10: 0-12-088782-7
For information on all Academic Press publications

visit our Web site at www.books.elsevier.com
Printed in the United States of America

06 07 08 09 10 9 8 7 6 5 4 3 2 1
Table of Contents
Part I Context 1
1. Introduction and Overview 3

1.1. Definition and introduction 4
1.2. History of research 7
1.3. Overview of the book 13
2. Importance and Roles of Soil Respiration 17

2.1. Soil respiration and ecosystem carbon balance 17
2.2. Soil respiration and nutrient cycling 21
2.3. Soil respiration and regional and global carbon cycling 22
2.4. Soil respiration and climate change 25
2.5. Soil respiration and carbon storage and trading 28
Part II Mechanisms 33
3. Processes of CO2 Production in Soil 35

3.1. Biochemistry of CO2 production processes 36
Tricarboxylic acid (TCA) cycle 36
Other CO2 production and consumption processes in soil 39
Respiratory quotient 40
vi Table of Contents
3.2. Root respiration 42

3.3. Rhizosphere respiration with labile carbon supply 46
3.4. Litter decomposition and soil organisms 49
3.5. Oxidation of soil organic matter (SOM) 55
4. Processes of CO2 Transport from Soil to the Atmosphere 61

4.1. CO2 transport within soil 61
4.2. CO2 release at the soil surface 67
4.3. CO2 transfer in plant canopy 70
4.4. CO2 transport in the planetary boundary layer (PBL) 74
Part III Regulation 77
5. Controlling Factors 79
5.1. Substrate supply and ecosystem productivity 79
5.2. Temperature 85
5.3. Soil moisture 92
5.4. Soil oxygen 98
5.5. Nitrogen 99
5.6. Soil texture 101
5.7. Soil pH 102
5.8. Interactions of multiple factors 104
6. Temporal and Spatial Variations in Soil Respiration 107

6.1. Temporal variation 108
Diurnal and weekly variation 108
Seasonal variation 110
Interannual variability 112
Decadal and centennial variation 113
6.2. Spatial patterns 115
Stand level 115
Landscape level 117
Regional scale 118
Biomes: Forests, grasslands, tundra, savannas/woodlands,
deserts, crop fields, and wetlands 120
6.3. Variation along gradients 128
Latitudes 128
Altitudes 129
Topography 130
Table of Contents vii
7. Responses to Disturbances 133
7.1. Elevated CO2 concentration 134
7.2. Climatic warming 138
7.3. Changes in precipitation frequency and intensity 143
7.4. Disturbances and manipulations of substrate supply 146
Fire or burning 146
Forest harvesting, thinning, and girdling 147
Grazing, clipping, and shading in grasslands 151
Litter removal and addition 152
7.5. Nitrogen deposition and fertilization 152
7.6. Agricultural cultivation 155
7.7. Interactive and relative effects of multiple factors 156
Part IV Approaches 159
8. Methods of Measurements and Estimations 161

8.1. Methodological challenges and classification of
measurement methods 162
8.2. Closed dynamic chamber (CDC) method 163
8.3. Open dynamic chamber (ODC) method 169
8.4. Closed static chamber (CSC) methods 170
Alkali trapping 171
Soda-lime trapping 172
8.5. Gas chromatograph (GC) 174
8.6. Chamber design and deployment 175
Chamber design 175
Chamber deployment 176
8.7. Gas-well (GW) method 178
8.8. Miscellaneous indirect methods 181
8.9. Method comparison 183
9. Separation of Source Components of Soil Respiration 187

9.1. Experimental manipulation methods 189
Direct component measurements and integration 189
Root exclusion 190
Severing substrate supply to the rhizosphere 190
Litter removal 194
9.2. Isotope methods 195
Growing C3 plants on C4 soil or C4 plants on C3 soil 197
CO2 enrichment experiments 199
viii Table of Contents
Bomb 14C tracer 204

Labeling experiments 207
9.3. Inference and modeling methods 209
Regression extrapolation and modeling analysis 209
Deconvolution analysis 210
9.4. Estimated relative contributions of different source
components 212
10. Modeling Synthesis and Analysis 215

10.1. Empirical models 216
Temperature-respiration models 216
Moisture-respiration models 219
Substrate-respiration models 224
Multifactor models 226
10.2. CO2 production models 230
10.3. CO2 production-transport models 239
10.4. Modeling soil respiration at different scales 241
10.5. Model development and evaluation 244
Appendix 247
References 257
Index 307
Preface
Soil respiration is an ecosystem process that releases carbon dioxide from

soil via root respiration, microbial decomposition of litter and soil organic
matter, and fauna respiration. Research on soil respiration has been remark-
ably active in the past decade partly because it is among the least understood
subjects in ecosystem ecology and partly because it represents the second
largest flux of carbon cycling between the atmosphere and terrestrial eco-
systems. As one key process of ecosystems, soil respiration is related to eco-
system productivity, soil fertility, and regional and global carbon cycles. Since
the global carbon cycle regulates climate change, soil respiration also be-
comes relevant to climate change, carbon trading, and environmental policy.
In short, soil respiration is nowadays a multidisciplinary subject that is of
concern not only to ecologists, soil scientists, microbiologists, and agrono-
mists but also to atmospheric scientists, biogeochemists, carbon traders, and
policy-makers. To date, no book has been published to synthesize extant
information on soil respiration in spite of its importance in many disciplines.
We write this book to fill this void and to stimulate broad interests in this
subject among students, scientists, environmental managers, and policy
makers from different disciplines,
x Preface
The active research in the past decade has substantially advanced our
understanding but, meanwhile, created much confusion with considerable
repetitive work in the research community. Much of the confusion and repeti-
tion stems from the lack of a systematic organization of knowledge on
fundamental processes of soil respiration. It was our initial motivation to lay
down the foundation of the soil respiration sciences and to clarify some of
the confusion. Toward that goal, we make an attempt to progressively intro-
duce and rigorously define concepts and basic processes. We also try to
structure the book in such a way that all the major up-to-dated research
findings can be logically summarized. The book is accordingly divided into
four sections—context, mechanisms, regulation, and approaches—and ten
chapters. Chapters 1 and 2 offer a contextual view of the soil respiration
science and lay down its relationships with a variety of issues in carbon
research. Chapters 3 and 4 describe fundamental processes of CO2 production
and transport. Chapters 5–7 present regulatory mechanisms of soil respira-
tion, including controlling factors, spatial and temporal variations, and
responses to natural and human-made perturbations. Chapters 8—10
illustrate research approaches to measurement of soil respiration, partition-
ing to various components, and modeling. It is our hope that this book helps
clarify confusion and identify knowledge gaps where research may be most
productive.
We write the book for undergraduate and graduate students, professors
and researchers in areas of ecology, soil science, biogeochemistry, earth
system science, atmosphere, climate molders, microbiology, agronomy, plant
physiology, global change biology, and environmental sciences. The book
introduces concepts and processes in a logical way so that students and
laymen who do not have much background in this area are can read the book
without too much difficulty. The book has also summarized the contempo-
rary research findings with extensive references. Scientists who are actively
working on soil respiration should find this book as a useful reference book
for their research. We also recognize that the field of soil respiration research
is evolving very quickly. Even within the time span from the manuscript
submission to the publication of this book, many important papers have been
published. Inevitably, many good papers may have been left out. We are sorry
if we miss your work in this book but welcome you to write us emails and
send us the postal mails with your important publications. We will try our
best to incorporate your work into the new version of the book in the
future.
This book is first dedicated to our fellow researchers. Their devotion to
and passionate on the soil respiration science are the impetus of advances in
our understanding on this subject. Their imagination and creativity result in,
for example, diverse ideas, experimental evidence from different angles, and
Preface xi
measurements by distinct methods. Their rigorous logic helps critique results,
identify new issues to be addressed, and generate new ideas to be tested. Their
meticulous methodology checks measurement and modeling results once and
again, enhancing the robustness of our knowledge. Their collective effort
helps establish the soil respiration science and, more importantly, bring it
into a focal research area in the earth system science. We hope that this book
will stimulate further interest in this fascinating subject and promote high-
quality scientific contribution.
We also dedicate this book to our families. Our parents taught us to work
hard no matter what we are doing, which becomes the lifetime gift to us. The
hardship of lives in our childhoods makes us appreciate what we have every-
day. We thank our spouses for their understanding of our career choices and
for their support to our effort on book writing. They have sacrificed countless
hours of family activities to make time for us to work on the book. Our chil-
dren brought us tremendous fun to our busy lives. In particular, Jessica Y.
Luo has read the first two chapters and offered suggestions to improve reader-
ship of the book.
Yiqi Luo is also grateful for students and post-doctoral fellows in his labo-
ratory who have worked with him to develop ideas, test various hypotheses,
and contribute to discussion in the research community via publications and
participation in international meetings.
Finally, we are indebted to many colleagues and authors who have sent us
reprints of their papers and manuscripts. We are grateful to Eric A. Davidson,
Joseph M. Craine, Dafeng Hui, Changhui Peng, Weixing Cheng, and Kiona
Ogle for their time to read the manuscript and for many helpful suggestions
and criticisms they have offered. We also thank Kelly D. Sonnack and Meg
Day of Academic Press/Elsevier for their patience and encouragement for this
project, Cate Barr for providing a cover design and Deborah Fogel for help in
editing manuscripts. Yiqi Luo thanks Dr. Lars Hedin for hosting his sabbati-
cal leave at Princeton University where the manuscript was finalized. Yiqi
Luo also acknowledges the financial support from US Department of Energy
and National Science Foundation, which has helped maintain his active
research in the past decade.
Yiqi Luo and Xuhui Zhou

Norman, Oklahoma
April 12, 2006
PART I
Context
CHAPTER 1
Introduction and Overview
1.1. Definition and introduction 4

1.2. History of research 7
1.3. Overview of the book 13
Soil respiration is a crucial piece of the puzzle that is the earth’s system. To
understand how the earth’s system functions, we need to figure out the role
that soil respiration plays in regulating atmospheric CO2 concentration and
climate dynamics. Will global warming instigate a positive feedback loop
between the global carbon cycle and climate system that would, in turn,
aggravate climatic warming? How critical is soil respiration in regulating this
positive feedback? To answer these questions, we have to understand the
processes involved in soil respiration, examine how these processes respond
to environmental change, and account for their spatial and temporal
variability.
Since climate change is one of the main challenges facing humanity, quan-
tification of soil respiration is no longer just a tedious academic issue. It is
also relevant to farmers, foresters, and government officials. Can respiratory
carbon emission and/or photosynthetic carbon uptake be manipulated to
maximize carbon storage so that farmers and foresters can earn cash awards
in global carbon-trading markets? To effectively manipulate respiratory
carbon emission from terrestrial ecosystems, we need to identify the major
factors that control soil respiration. Even if we can manipulate respiratory
processes, how could signatory countries to the Kyoto treaty verify carbon
sinks in the biosphere to claim their credits during the intergovernmental
negotiations? All these issues make it necessary for us to invent reliable
methods to measure soil respiration accurately in croplands, forest areas, and
other regions. Can the managed carbon sinks last long enough to mitigate
greenhouse gas emission effectively in the future? How will soil respiration
respond to natural and human-made perturbations? To answer all these ques-
tions, it is necessary to develop a predictive understanding of soil respiration,
aiming toward a mechanistic modeling of soil respiration. It is evident from
all these examples that studying soil respiration is not only desirable for
3
4 Chapter 1 Introduction and Overview
200
Number of papers published on soil respiration
150
100
50
0
1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004
Years
FIGURE 1.1 Number of papers published on soil respiration since 1985. The number was
obtained from a search for the key terms “soil respiration,” “soil CO2 efflux”, and “belowground
respiration” in the Web of Science database.
purely academic reasons but also crucial in the commercial and political
arenas.
Due to the recent societal need to mitigate climate change and the scientific
aspiration to understand soil respiration itself, the research community has
been very active in studying soil respiration. During the past 15 years, the
number of papers published on soil respiration has linearly increased and
reached nearly 200 papers in 2003–2004, compared with about 10 papers in
1985–1990 (Fig. 1.1). The active research also partially reflects the fact that
soil respiration remains least understood among ecosystem carbon processes,
despite its central role in the global carbon cycle and climate change. This
book lays down the fundamentals of soil respiration while synthesizing the
recent literature in this field.
1.1. DEFINITION AND INTRODUCTION
The word respiration, derived from the Latin prefi x re- (back, again) and root
word spirare (to breathe), literally means breathing again and again. It is thus
used to describe the process of gas exchange between organism and environ-
ment. Physiologically, respiration is a series of metabolic processes that break
down (or catabolize) organic molecules to liberate energy, water, and carbon
Definition and Introduction 5
dioxide (CO2) in a cell. All living organisms—plants, animals, and micro-
organisms alike—share similar pathways of respiration to obtain the energy
that fuels life while releasing CO2. Respiration is often studied in relation to
energy supply at the biochemical and cellular levels as a major component of
bioenergetics. However, bioenergetics in soils is not well developed (Dilly
2005), and soil respiration is studied predominantly in relation to CO2 and
O2 exchanges. In this book the word respiration is used mainly to describe
CO2 production rather than energy supply.
For the purposes of this book, soil respiration is defined as the production
of carbon dioxide by organisms and the plant parts in soil. These organisms
are soil microbes and fauna, and the plant parts are roots and rhizomes in
the soil. Additionally, soil is often defined as a mixture of dead organic matter,
air, water, and weathered rock that supports plant growth (Buscot 2005).
Some authors (e.g., Killham 1994) also include living organisms in the defini-
tion of soil, treating roots, soil microbes, and soil fauna as part of soil. There-
fore, it makes sense to talk about soil that can breathe. Soil respiration means
that the living biomass of soil respires CO2, while soil organisms gain energy
from catabolizing organic matter to support life.
Soil respiration is sometimes called belowground respiration, in contrast
with aboveground respiration. The latter refers to respiratory CO2 production
by the plant parts above the soil surface. Although the definition of soil usually
does not include dead plant materials at the soil surface that have not been
well decomposed, CO2 production via litter decomposition in the litter layers
is generally included in soil respiration (or belowground respiration) in many
publications and, for the sake of simplicity, in this book as well.
Technically, the rate of CO2 production in the soil (i.e., the soil respiration
rate) cannot be directly measured in the field. Measurements are often made
at the soil surface to quantify a rate of CO2 efflux from the soil to the atmos-
phere. The instantaneous rate of soil CO2 efflux is controlled not only by the
rate of soil respiration but also by the transport of CO2 along the soil profile
and at the soil surface (see Chapter 4). The CO2 transport is influenced by
the strength of the CO2 concentration gradient between the soil and the
atmosphere, soil porosity, wind speed, and other factors. At a steady state,
the CO2 efflux rate at the soil surface equals the rate of CO2 production in
soil. In this case, soil CO2 efflux is practically equivalent to soil respiration,
and the two terms are thus interchangeable.
However, there are several situations in which CO2 production may not be
at a steady state with CO2 transport. For example, soil degassing occurs
during rainfall or irrigation, driving CO2 stored in the soil air space out of
the soil. After rainfall or irrigation, CO2 produced by soil organisms is par-
tially stored in the soil to rebuild the CO2 concentration gradient. Carbonic
acid reaction and microbial methanogensis could each produce or consume
CO2, depending on conditions that influence reaction equilibriums (see

Chapter 3). Thus, the CO2 released at the soil surface could be generated by
carbonic acid reactions during rock weathering, particularly in arid lands
where carbonic reaction is very strong. On the other hand, the CO2 produced
by soil living tissues could be absorbed by microbes during methanogenic
processes. However, the amount of CO2 produced and/or consumed by car-
bonation and methanogenesis is generally trivial in comparison with soil
respiration, except in very dry lands. The non-steady-state CO2 efflux at the
soil surface occurs mostly during rainfall or irrigation after long periods of
drought (Liu et al. 2002a, Xu et al. 2004). In absence of major perturbation,
the rate of CO2 production in soil is indistinguishable from the rate of
CO2 efflux at the soil surface on a daily or longer time-scale (Hui and Luo
2004). Thus, the term soil respiration is practically interchangeable with
soil surface CO2 efflux on a long-term scale. However, soil CO2 efflux rates
measured at shorter time-scales may not be equivalent to the rate of soil
respiration.
Soil respiration usually accounts for the majority of ecosystem respiration,
which is the sum of soil respiration and respiration of aboveground parts of
plants (see Chapter 2). Some methods can directly measure ecosystem respi-
ration, from which soil respiration is estimated indirectly (see Chapter 8).
Thus, the soil and ecosystem respirations are closely related. Although this
book focuses on soil respiration, it often describes ecosystem respiration as
well.
As a preview, Figure 1.2 shows a typical time course of CO2 efflux rates
from soil. The time course, which was measured at the soil surface in a tall-
grass prairie of Oklahoma, displays a distinct seasonal pattern of high soil
respiration during summer and low respiration in winter. The seasonal
pattern is roughly repeated in subsequent years. Nonetheless, there are
observable variations from year to year. For example, the summer peak of
soil respiration reaches nearly 6 µmol m−2 s −1 in 2002 and is less than 4 µmol
m−2 s −1 in 2001. The winter low is nearly 0 µmol m−2 s −1 in 2002 but 0.3–
0.5 µmol m−2 s −1 in other years. In most years, there are dips in the measured
soil respiration during the late summer and early autumn, but in 2004 the
seasonal pattern is relatively smooth. This kind of year-to-year variation
exemplifies the term “interannual variability.”
Similar seasonal patterns have also been observed in northern semiarid
grasslands (Frank et al. 2002), forests (Salvage and Davidson 2001, Epron et al.
2004, King et al. 2004), and croplands (Beyer 1991). For example, soil respira-
tion varies from nearly 0 µmol m−2 s−1 in the winter to about 10 µmol m−2 s−1 in
the summer over one year in the Duke Forest, North Carolina (King et al. 2004).
This seasonal pattern repeats from 1997 to 2002, and interannual variation is
apparent with different peaks in summer and valleys in winter.
History of Research 7
Soil respiration (µmol m -2 s-1) 6
0
1999 2000 2001 2002 2003 2004 2005
Year
FIGURE 1.2 Measured rate of soil CO2 efflux in a tallgrass prairie of Oklahoma, USA from
1999 to 2005. Open circles represent data points, and bars indicate the one standard error
below and above the data points. Data are only for the measured soil CO2 efflux in the control
treatment in a warming and clipping experiment and adopted from Luo et al. (2001), Wan
et al. (2005), and Zhou et al. (2006).
From the observed soil respiration patterns, we can ask many questions.
For example, what causes such seasonal and interannual variations? Why
does soil respiration vary from one site to another? How can we scale up the
plot-level measurements to estimate total carbon losses on regional and global
scales? Can we derive general mechanisms from the observed patterns and
then predict future changes in soil respiration? What percentage of the lost
carbon is from root respiration? How much is the carbon released by soil
respiration directly from the recent photosynthesis? This book will address
these questions, among others, as it lays down the basic principles of soil
respiration. Before turning to these issues, however, let’s first review the
history of research on soil respiration.
1.2. HISTORY OF RESEARCH
Research on soil respiration has an impressively long history (Fig. 1.3) and
can be dated back to papers by Wollny (1831), Boussingault and Levy (1853),
and Möller (1879). The earliest studies of soil respiration were intended to
characterize soil metabolism. Twentieth-century research on soil respiration
can be divided into roughly four major periods. During the first few decades
of the century, research on soil respiration was conducted primarily in the
laboratory with agricultural soil. Soil respiration was used to evaluate soil
fertility and biological activities in soil. Chemical fertilizers, invented in the
late 19th century, were applied to crops to stimulate growth and considerably
enhanced agricultural productivity as a result. At that time, researchers
emphasized understanding the soil properties that influence crop production.
Soil respiration was used as an index of soil fertility for agricultural produc-
tion (Russell and Appleyard 1915), because in a field study, fertilization of
agricultural crops generally increases soil respiration rates (Lundegårdh
1927). Some laboratory studies, however, showed that nutrient release was
not proportional to the carbon release during mineralization (Waksman and
Starkey 1924, Pinck et al. 1950).
During that period, some primitive methods for the measurement of soil
respiration were developed. Stoklasa and Ernest (1905) passed CO2-free air
over soil samples contained in a flask and measured the amount of CO2
released from the soil samples using the alkali absorption method.
Lundegårdh (1927) recognized that measured CO2 efflux from soil samples
in the laboratory might not be representative of that from intact soils in the
field, where, he argued, diffusion was a chief process controlling efflux of
CO2. He was probably the first scientist to make in situ measurements of rates
of CO2 efflux from field soil by covering the soil surface with a chamber for
a period of time. Then he took air samples with brass tubes from the chamber,
as well as from air spaces in the soil at three different depths. The air samples
were passed through alkali solutions for measurements of soil respiration.
Humfeld (1930) modified Lundegårdh’s method and passed air through the
chamber with inlet and outlet ports to collect the CO2-enriched air in an
alkali absorption train. The alkali absorption chamber method, first intro-
duced by Lundegårdh (1921), modified by Humfeld (1930) and others, and
widely used in the following decades, places static alkali solution within the
chamber followed by titration of chloric acid.
By this time the major factors that influence soil respiration had been
identified. Greaves and Carter (1920) were among the first to document a
consistent relationship between soil water content and microbial activity.
Turpin (1920) reviewed soil respiration and concluded that the primary
source of CO2 efflux from soils was attributable to bacterial decomposition.
Lundegårdh (1927) pointed out that soil diffusion was important in control-
ling the efflux of CO2. Smith and Brown (1933) indicated that the rate of dif-
fusion of CO2 through the soil correlated with CO2 production. Lebedjantzev
(1924) observed that air drying of soil samples increased fertility (such as
NH4-N, amide-N, and phosphorus) of a variety of soils and decreased the
number of microorganisms in pot experiments.
Few publications on soil respiration can be identified during the relatively
inactive research period from the late 1930s to the early 1950s, possibly due
to the worldwide social turbulence of that period. From the late 1950s to the
1970s, research activity on soil respiration resumed (Fig. 1.3), mainly from
an ecological perspective, as scientists tried to understand heterotrophic
processes in the soils of native ecosystems (Lieth and Ouellette 1962, Witkamp
1966, Raguotis 1967, Schulze 1967, Reiners 1968, Kucera and Kirkham 1971).
During that period, research advanced the science of soil respiration in many
respects, including (1) methods of measurement, (2) controlling factors, (3)
partitioning into components, (4) relationships with other ecosystem carbon
processes, and (5) synthesis and scaling to global estimation.
Many studies were devoted to careful evaluation of the various factors that
affect the accuracy of the alkali absorption method (Walter 1952, Howard
1966, Kirita and Hozumi 1966, Kirita 1971, Chapman 1971, 1979, Anderson
1973, Gupta and Singh 1977). The accuracy of the method was found to vary
with factors such as the amount and strength of alkali used, the area of
covered soil, the chamber height above the ground, the depth of the chamber
inserted into soil, the surface area and the height of the alkali container
within the chamber, the duration of measurement, and the rates of soil CO2
efflux. Minderman and Vulto (1973) suggested the use of fine-grained soda
lime instead of alkali solution to absorb CO2.
Closed static chamber method with Alkali method & Gas well method
Dynamic chamber methods with IRGA
Method
Micrometeorological techniques
Soil metabolism Soil fertility Ecosystems No tillage Global change
1850 1900 1950 2000
C trading
Issues
Climate feedback
Soil metabolism
Global C cycle
Nutrient release Annual carbon balance
Soil fertility Component partitioning
Regulatory factors: temperature & moisture
FIGURE 1.3 Schematic illustration of the history of soil respiration research since the 1830s.
Within the main axis are major themes in different eras of research. There is little research
activity from late 1930s to early 1950s. Above the axis is method development for measurement
of soil respiration. Below the axis are major issues that have been addressed by and/or motivate
soil respiration research during the different eras.
One major technical advance was made in the 1950s: infrared gas analyzer
(IRGA) was used for the measurement of soil respiration. Haber (1958) first
used IRGA to calibrate the alkali absorption method. Golley et al. (1962) were
among the first to make field measurements of soil respiration on the peat
floor of a mangrove forest using IRGA. Reiners (1968) examined how gas flow
rates influenced IRGA measurement of CO2 evolution, while Kanemasu et al.
(1974) studied effects of air “suction” and “pressure” on IRGA measurements
of soil respiration. Measured CO2 efflux with the suction chamber was one
order of magnitude higher than with the pressure chamber. The suction
chamber drew CO2 from the soil outside the chamber and/or in deep layers
via mass flow. Edwards and Solins (1973) designed an open flow system with
the chamber linked to IRGA to measure soil respiration continuously. Edwards
(1974) used movable chambers that were lowered onto the forest floor during
measurements and lifted between measurements. The movable chambers
allowed natural drying of the soil and litterfall onto the measurement surface.
The IRGA measurements of soil CO2 efflux were then compared with those
using the alkali absorption method (Kirita and Hozumi 1966). Many studies
found that the alkali method underestimated soil CO2 efflux compared with
the IRGA measurements (Haber 1958, Witkamp 1966, Kucera and Kirkham
1971). Other studies did not detect any significant differences between the
two methods (e.g., Ino and Monsi 1969).
The gas-well method first used by Lundegårdh (1927) to estimate soil
respiration from a CO2 concentration gradient along soil profiles was fully
developed by de Jong et al. (1979). Meanwhile, a variety of micrometeorologi-
cal methods, such as Bowen ratio and eddy flux, have been developed to
measure gas exchanges within and above the plant canopy (Monteith 1962,
Monteith et al. 1964), from which soil respiration was indirectly estimated.
From the late 1950s to the 1970s, knowledge of factors that regulate soil
respiration was greatly enriched. Bunt and Rovira (1954) studied soil respira-
tion in a temperature range of 10 to 70°C. They found that O2 uptake and
CO2 release increased with temperature up to 50°C, above which it declined.
Many studies demonstrated that soil respiration correlated exponentially
with temperature (Wiant 1967, Kucera and Kirkhma 1971, Medina and Zelwer
1972). Drobnik (1962) estimated Q10, that is, a quotient indicating the tem-
perature sensitivity of soil respiration (see Chapter 5), to be 1.6 to 2.0 in
response to temperatures ranging from 8 to 28°C. Wiant (1967) estimated
Q10 to be approximately 2 for temperatures from 20 to 40°C. Soil moisture
was also identified as important in influencing soil respiration. A laboratory
study suggested that microbial respiration decreased when soil moisture was
below 40% or above 80% of the field-holding capacity (Ino and Monsi 1969).
Soil temperature and moisture combined could account for up to 90% of the
variation of soil respiration measured in the field (Reiners 1968).
Birch and his colleague (Birch and Friend 1956, Birch 1958) conducted a
notable study demonstrating that when a soil was dried and rewetted, decom-
position of its organic matter was enhanced, leading to a flush of CO2 produc-
tion. They explained that the drying-wetting effect was not related to microbial
stimulation or microbial death but rather caused by liberation of rapidly
decomposable material from the clay. The clay protected the organic materials
from microbial attacks under consistently moist conditions.
During that period, components of soil respiration were clearly identified
into two major categories: autotrophic and heterotrophic respiration. The
autotrophic components are the metabolic respiration of live root, associated
mycorrhiza, and symbiotic N fi xing nodules. The heterotrophic respiration
is from microbial decomposition of root exudates in rhizosphere, above-
ground and belowground litter, and soil organic matter (SOM). Coleman
(1973b) measured total respiration of intact soil cores and individual compo-
nents of roots, litter, and soil. Contribution to the total soil respiration was
8 to 17% from roots, 6 to 16% from litter, and 67 to 80% from soil microbes
in a successional grassland. Edwards and Sollins (1973) partitioned total soil
respiration from a forest into 35% from roots, 48% from litter, and 17% from
soil. Richards (1974) found it difficult to partition soil respiration among dif-
ferent soil fauna, fungi, and bacteria.
Field measurements over the whole growing seasons made it possible to
scale up individual measurements to estimate annual carbon efflux. Kucera
and Kirkham (1971) estimated annual soil CO2 efflux to be 452 g C m−2 yr−1 in
a tallgrass prairie by applying a temperature-respiration regression to continu-
ous temperature records. Coleman (1973a) scaled up monthly averages of soil
respiration in a grassland and estimated annual soil CO2 efflux to be 357 to
421 g C m−2 yr−1. Estimated annual soil CO2 releases were about 1000 g C m−2 yr−1
in many forests (Edwards and Sollins 1973, Garrett and Cox 1973).
Estimated annual efflux from soil respiration was often compared with
annual carbon influx via aboveground litterfall, although the two processes
are not completely comparable. Reiners (1968) showed that total soil respira-
tory carbon release was three times higher than litter carbon input. Edwards
and Sollins (1973) found that litter decomposition accounted for only one-
fifth of annual soil respiration. Anderson (1973) showed that annual soil
respiration released 2.5 times as much carbon in annual litterfall. However,
several studies demonstrated that carbon released by soil respiration was
equivalent to that input from litterfall (Colemen 1973a, Witkamp and Frank
1969).
The accumulation of studies during that period offered opportunities to
synthesize and compile results from many ecosystems. Singh and Gupta
(1977) produced a major synthesis on the carbon processes of litter decom-
position, soil respiration, root respiration, microbial respiration, faunal
respiration, and SOM dynamics. Schlesinger (1977) reviewed many studies

on soil respiration in the literature in order to develop latitudinal patterns of
soil respiration worldwide and estimate a global total of carbon released via
soil respiration.
Bunnell et al. (1977) and Minderman (1968) suggested that decomposition
could best be represented by the summation of the exponential decay curves
for all major chemical constituents, including sugars, cellulose, hemicellu-
lose, lignin, waxes, and phenols. Henin et al. (1959) appeared to have been
the first to propose a model that explicitly relates the two exponential rates
to fresh plant carbon and “humified” carbon.
Long-term no-till plots were first established at the International Institute
of Tropical Agriculture, Ibadan, in 1971 and continued through 1987 (Lal
2004). In the 1980s the agricultural practice of no tillage stimulated research
on soil properties. Soil respiration was often used to indicate biological activi-
ties in soil with different tillage treatments (Anderson 1982). For example,
Linn and Doran (1984) studied how no tillage affected soil water–filled pore
space and its relationships with CO2 and N2O production. The level of soil
aeration using microbial respiration rates of aerobic heterotrophs was also
examined for compaction problems in a no-tillage management system (Linn
and Doran 1984, Wilson et al. 1985, Neilson and Pepper 1990).
Since the 1990s, research on soil respiration has been driven primarily by
global change. While climate research has its own long history (Weart 2003),
the ecology research community, stimulated by the International Geosphere
Biosphere Program (IGBP) and by a U.S. National Research Council (NRC)
report (NRC 1986), has been involved in global change research in the past
two decades and has studied ecosystem-level responses to climate change
since the early 1990s (Mooney et al. 1991). In particular, the paper by Tan et
al. (1990) played a critical role in attracting researchers’ attention to the land
biosphere. Their analysis of atmospheric CO2 data suggested that land bio-
sphere may absorb a large portion of the emitted carbon from anthropogenic
sources. Three reports by the Intergovermental Panel on Climate Change
(IPCC, 1990, 1995, 2001) and Schimel (1995) provided a global perspective
on the carbon cycle in terrestrial ecosystems. Cox et al. (2000) linked a
carbon cycle model with a global circulation model and highlighted the
importance of the temperature sensitivity of respiration in future climatic
predictions. That study continues to stimulate great interest in the tempera-
ture sensitivity of soil respiration among the research community.
Advances in measurement techniques have also stimulated modern, active
research on soil respiration. Portable IRGAs have been widely used
to measure soil surface CO2 fluxes since the early 1990s (Norman et al.
1992). The IRGA method requires relatively less technique training than the
traditional alkali or soda-lime absorption methods, but it provides quicker
Overview of the Book 13
measurements of soil surface CO2 effluxes. Meanwhile, many companies have
retooled IRGA sensors and developed various chambers specifically for the
measurement of soil CO2 effluxes (see Chapter 8 and Appendix) facilitating
research on soil respiration.
1.3. OVERVIEW OF THE BOOK
This book, which comprises 10 chapters, is dedicated to providing an under-

standing of various aspects of soil respiration. Chapters 1 and 2 provide a
context of soil respiration science. Chapters 3 and 4 describe fundamental
processes of CO2 production and CO2 transport. Chapters 5 through 7 present
regulatory mechanisms of soil respiration, including controlling factors, spatial
and temporal variations, and responses to natural and human-made perturba-
tions. Chapters 8 through 10 discuss research approaches to measurement of
soil respiration, partitioning to various components, and modeling.
Following the introduction and brief history of research on soil respiration
covered in this chapter, Chapter 2 places soil respiration in the context of
ecosystem carbon balance, nutrient cycling, regional and global carbon
cycling, climate change, and carbon storage and trading. Soil respiration
releases a large portion of carbon fi xed by photosynthesis and strongly regu-
lates net ecosystem productivity. Carbon dioxide released via microbial
decomposition of litter and SOM is accompanied by either immobilization or
mineralization of nutrients and is thus related to soil nutrient dynamics. Soil
respiration plays a critical role in regulating global and regional carbon
cycles. Its temperature sensitivity is a key issue in modeling feedback between
global carbon cycling and climate change in response to anthropogenic
warming. Although it is not the direct mechanism underlying land carbon
storage, soil respiration is relevant to understanding carbon sequestration and
global carbon trading markets.
Chapter 3 focuses on the processes of CO2 production, including the fun-
damental biochemistry of respiratory processes, root respiration, microbial
respiration in rhizosphere, and microbial decomposition of litter and SOM.
The primary biochemical process of CO2 production is the tricarboxylic acid
(TCA) cycle. Root respiration in an ecosystem is determined by root biomass
growth and the specific rates of root respiration. Microbial respiration occurs
while root exudates are broken down, litter is decomposited, and SOM oxi-
dated. Microorganism communities that use root exudates, litter, and SOM
as substrates differ greatly and are briefly described in this chapter.
Chapter 4 describes processes of CO2 transport along vertical pro-
files within the soil, at the soil surface, in the canopy, and in the planetary
boundary layer. Soil CO2 transport is driven primarily by gradients of CO2
concentration along soil vertical profiles and determined by diffusion and

mass flow processes. The CO2 release at the soil surface depends on CO2
gradients and is strongly affected by wind gusts, turbulences, and atmos-
pheric pressure fluctuation. The CO2 transport in the canopy and planetary
boundary layer may not be directly relevant to soil respiration per se but is
influenced by and often used to estimate soil respiration indirectly.
Soil respiration is affected by many factors, such as substrate supply, tem-
perature, moisture, oxygen, nitrogen, soil texture, and pH value. Chapter 5
focuses on how individual factors regulate component processes of soil res-
piration and attempts to show that many of the factors influence multiple
processes in various magnitudes and at different directions, leading to
variable responses and complex patterns of soil respiration. The interactive
effects of multiple factors on soil respiration are very complex and poorly
understood.
Chapter 6 presents spatial and temporal patterns of soil respiration. It
discusses temporal variations in soil respiration at multiple time-scales—
from diurnal and weekly to seasonal, interannual, and decadal and centen-
nial. Spatial patterns emerge at the stand level, landscape and regional scales,
and across biomes. The chapter comparatively presents soil respiration among
ecosystem types and examines general relationships of soil respiration to
ecosystem productivities, prevailing environmental variables, and soil
characteristics. This chapter also examines how soil respiration varies along
latitudinal, altitudinal, and topographical gradients.
Chapter 7 describes changes in soil respiration in response to a variety of
perturbations, such as elevated CO2, global warming, changes in precipitation
frequency and intensity, disturbances and manipulation of substrate supply,
nitrogen deposition and fertilization, and agricultural cultivation. Generally
speaking, soil respiration increases when substrate availability increases,
such as under elevated CO2 and litter addition. Soil respiration decreases if
substrate supply is reduced under disturbances of fire, burning, forest cutting,
cutting and grazing in grasslands, and litter removal. Agricultural cultivation
usually stimulates soil respiration in the short term because of soil distur-
bances but results in a long-term decrease in soil carbon content. Climatic
warming also causes short-term stimulation of soil respiration and may
induce long-term acclimation. Responses of soil respiration to changes in
precipitation and nitrogen addition are highly variable.
Chapter 8 introduces a variety of methods for measurement of soil respira-
tion. The most commonly used are chamber methods, which include the
closed dynamic-chamber method, the open dynamic-chamber method, and
the closed static-chamber method. Soil respiration can also be estimated from
air samples from different depths of soil using the gas-well method. This
chapter describes the basic principles behind those methods, discusses
Overview of the Book 15
chamber designs and deployment, and assesses the accuracy and potential
issues of those methods. It also briefly describes a few indirect methods for
estimation of soil respiration.
The partitioning of soil respiration is critical for developing predictive
understanding of soil respiration. Chapter 9 introduces three groups of
methods—experimental manipulations, isotope tracers, and indirect infer-
ence analysis—for partitioning. The experimental methods manipulate the
substrate supply to different pathways of soil respiration and separate com-
ponents of soil respiration. The isotope methods take advantages of isotope
signals of C3 and C4 plants and soils, CO2 experiments that fumigate CO2
with different isotope values, bomb 14C that enriched 14C in the atmosphere
in 1950s and 1960s, and labeling experiments. The inference methods are to
estimate component contributions through regression extrapolation and
deconvolution analysis. This chapter summarizes estimates of contributions
of each source component to the total soil respiration.
Chapter 10 provides a general description of models and modeling studies
of soil respiration. In general, the modeling studies are based on three types
of models: empirical models, CO2 production models, and CO2 production
and transport models. The empirical models are derived primarily from
regression analysis of soil respiration with temperature, moisture, and some
surrogate quantities of substrate availability. The production models usually
incorporate carbon processes of photosynthesis, partitioning, and decompo-
sition of litter and SOM. The production-transport models consider transport
processes of soil CO2 along a soil profile from the production sites to soil
surface. This chapter examines modeling studies according to different spatial
and temporal scales and discusses model development and evaluation.
CHAPTER 2
Importance and Roles of
Soil Respiration
2.1. Soil respiration and ecosystem carbon

balance 17
2.2. Soil respiration and nutrient cycling 21
2.3. Soil respiration and regional and global
carbon cycling 22
2.4. Soil respiration and climate change 25
2.5. Soil respiration and carbon storage and
trading 28
Soil respiration is a subject that is of concern not only to ecologists but also
to scientists who study atmospheric dynamics and earth system functioning.
As an integral part of the ecosystem carbon cycle, soil respiration is related
to various components of ecosystem production. Soil respiration is also
intimately associated with nutrient processes such as decomposition and
mineralization. Moreover, soil respiration plays a critical role in regulating
atmospheric CO2 concentration and climate dynamics in the earth system.
Thus, it becomes relevant to the mitigation of climate change and the imple-
mentation of international climate treaties in terms of carbon storage and
trading. This chapter relates soil respiration to ecosystem carbon balance
and production, nutrient cycling, regional and global carbon cycling, climate
change, and carbon storage and trading.
2.1. SOIL RESPIRATION AND ECOSYSTEM

CARBON BALANCE
The carbon cycle in an ecosystem usually initiates when plants fix CO2 from
the air and convert it to organic carbon compounds through photosynthesis
(Fig. 2.1). Some of the organic carbon compounds are used to grow plant
tissues. Some are broken down to supply the plants with energy. During this
17
18 Chapter 2 Importance and Roles of Soil Respiration
CO2 NEE
Photosynthesis Ecosystom respiration Re
GPP
Ra Soil respiration Rs
Rm
Rb
Root SOM Litter Decomposition
Rp=Ra–Rb
Leaching losses
NPP=GPP-Rp
FIGURE 2.1 Schematic diagram of ecosystem carbon processes. Abbreviation see text.
process, CO2 is released back into the atmosphere through plant respiration.
The grown tissues include leaves, stems (e.g., wood for trees), and roots.
Leaves and fine roots usually live for several months up to a few years before
death, whereas woody tissues may grow for hundreds of years in forests. Dead
plant materials (i.e., litter) are decomposed by microorganisms to provide
energy for microbial biomass growth and other activities. At the same time,
CO2 is released back into the atmosphere through microbial respiration. The
live microbial biomass is mixed with organic residuals of dead plants and
dead microbes to form soil organic matter (SOM). SOM can store carbon in
soil for hundreds and thousands of years before it is broken down to CO2
through respiration by microbes.
Through the carbon cycle, CO2 is produced by both plant respiration (Rp)
and microbial respiration (Rm) that occurs during decomposition of litter and
SOM. Rp is often called autotrophic respiration and can be separated into
aboveground plant respiration (Ra) and belowground plant respiration (Rb).
(The belowground plant respiration is often equivalent to root respiration.)
Microbial respiration (Rm) during the decomposition of litter and SOM is
called heterotrophic respiration. The efflux rate measured at the soil surface
(Rs) is the sum of root respiration and microbial respiration:
R s = R b + Rm (2.1)
The CO2 efflux measured at the soil surface can be considered as soil
respiration when CO2 production and transport are at a steady state (see
Soil Respiration and Ecosystem Carbon Balance 19
Chapter 1). Thus, ecosystem respiration (Re), the total CO2 emission from an
ecosystem, can be estimated by:
Re = R a + R s (2.2)
The relationship of Rs with Re, as seen in equation 2.2, is well illustrated
by data collected from an aspen-dominated mixed hardwood forest in
Michigan from 1999 to 2003 (Curtis et al. 2005). On average, over the five
years Rs accounts for 71% of Re, while leaves and aboveground live wood
combined (Ra) contribute the rest of Re (Table 2.1). The relative contribution
of Rs to Re varies considerably in a year. Rs contributes nearly 100% of Re for
most of the winter; the contribution drops to about 60% during the period
of fast leaf expansion and then gradually increases during the growing season
as soil warms, reaching about 75% at the time of leaf abscission in the autumn
(Curtis et al. 2005). Typically, Rs contributes 30–80% of Re in forests.
Soil respiration is not only an important component of ecosystem respira-
tion but also closely related to ecosystem production such as gross primary
production (GPP), net primary production (NPP), and net ecosystem produc-
tion (NEP). GPP is annual carbon assimilation by photosynthesis ignoring
photorespiration. In the Michigan forest, for example, soil respiration is
approximately 63% of GPP (Table 2.1). NEP is GPP minus Re and also related
to soil respiration by:
NEP = GPP − Ra − Rs (2.3a)
or Rs is related to NEP though NPP, which is GPP minus autotrophic plant
respiration, by:
NEP = NPP − Rm
(2.3b)
= NPP + Rb − Rs
TABLE 2.1 Various components of ecosystem carbon fluxes in a mixed hardwood forest
from 1999 to 2003
Year Rs Re R s/Re GPP* R s/GPP NPP NEP
1999 1116 1538 0.73 1637 0.68 656 99

2000 987 1396 0.71 1580 0.62 678 184
2001 1005 1412 0.71 1615 0.62 704 203
2002 946 1404 0.67 1549 0.61 618 145
2003 960 1375 0.70 1545 0.62 650 170
Mean 1003 1425 0.71 1585 0.63 661 160
Note: GPP was estimated by a biometrical approach that sums up different components. The
biometrically estimated GPP was higher than that estimated by eddy-flux measurements by
nearly 30%. Units are g C m−2 yr−2. Modified with permission from New Phytologist: Curtis
et al. (2005)
Equation 2.3 is a quantitative basis of the biometrical approach to estima-

tion of net carbon storage in an ecosystem (i.e., NEP). NPP can be estimated
by measuring yearly increments in plant biomass. Ra is often estimated from
measured respiration rates of aboveground plant parts (i.e., leaves and live
wood in forest). Rb is estimated either from measured respiration rates of roots
or indirectly from Rs through partitioning techniques (see Chapter 9). With
measured soil respiration, NEP can be estimated from Equation 2.3. In the
Michigan hardwood forest, the estimated NEP by the biometrical method
ranged from 100 to 200 g C m−2 yr−1 (Table 2.1) (Curtis et al. 2005).
Another rate of flux in the ecosystem carbon cycle that can be relatively
easily measured, especially in forests, is aboveground litterfall. For a long
time scientists have sought a relationship between measured litterfall and soil
respiration (e.g., Reiners 1968). By synthesizing experimental results from
many forests in different regions with various types and ages of forests, Raich
and Naderhoffer (1989) generalized the relationship (Fig. 2.2) as:
Rs = aLa + b (2.4)
where La is aboveground litterfall and a and b are coefficients. Both Rs and La
are expressed in units of g C m−2 yr−1. The regression coefficient a is usually
about 3 (Raich and Naderhoffer 1989, Davidson et al. 2002a), suggesting that
carbon release from soil respiration is nearly three times the carbon input
2000
1800
Soil respiration (g C m-2 yr-1)
1600
1400
1200
1000
800
600
400
200
0
0 100 200 300 400 500 600
Litterfall (g C m-2 yr-1)

FIGURE 2.2 Correlation of soil respiration with the amount of aboveground litterfall across
many forest ecosystems (Redrawn with permission from Ecology: Raich and Naderhoffer
1989).
Soil Respiration and Nutrient Cycling 21
TABLE 2.2 Annual carbon fluxes for mid-rotation loblolly pine plantations
Fertilized and
Component Control Irrigated Fertilized Irrigated
Soil CO2 release 1263 1489 1293 1576

Root respiration 663 745 942 1062
Microbial respiration 600 744 351 514
NPP 500 635 1020 1235
NEP −100 −109 669 721
Note: Units are g C m−2 yr−1. Modified with permission from Canadian Journal of Forest Research
Maier and Kress (2000).
from aboveground litter. Indeed, soil respiration releases carbon from sources
of root litter, root exudates, and root respiration in addition to the above-
ground litterfall. The correlation was poor, however, among years at a single
site (Davidson et al. 2002a).
The relationships of Rs with other fluxes can also be used to examine
responses of an ecosystem to perturbations. Table 2.2, for example, presents
annual carbon fluxes in mid-rotation loblolly pine plantations as affected by
fertilization and irrigation (Maier and Kress 2000). Annual Rs is mainly
affected by irrigation, ranging from 1263 to 1576 g C m−2 yr−1 among the four
treatments. Belowground root respiration (Rb) is much more responsive to
fertilization than to irrigation, whereas Rm is considerably depressed by fer-
tilization. As a consequence, the relative contribution of Rb to Rs increases
from 52% under control to 73% under fertilization. Fertilization substantially
increased NPP, resulting in net carbon storage in the forest. NEP is negative
by 100 g C m−2 yr−1 without fertilization and becomes positive to 700 g C m−2 yr−1
with fertilization (Table 2.2).
2.2. SOIL RESPIRATION AND NUTRIENT CYCLING
A major component of soil respiration is from microbial decomposition of

litter and SOM that releases CO2, meanwhile immobilizing or mineralizing
nutrients (Coleman et al. 2004). During the initial phases of decomposition,
nitrogen that is mineralized from litter substrate is simultaneously immobi-
lized by microbes for their own growth, leading to an increased nitrogen
concentration in the mixture of litter substrate and microbes. Since the litter
substrate and microbes are not easily separated, in practice the mixture is
also called litter. The nitrogen concentration of decomposing litter usually
increases, while the absolute amount of nitrogen in the litter may or may not
increase during the decomposition. The absolute amount of nitrogen increases

when nitrogen from exogenous sources in soil or from fi xation is incorporated
into microbial biomass growth. The release of carbon combined with nitrogen
immobilization during the litter decomposition gradually decreases carbon-
nitrogen ratio (C : N) until mineralized nitrogen from litter substrate is greater
than required for microbial growth. After that point, litter decomposition
leads to a net release of nitrogen. Similarly, phosphorus and sulfur may also
increase in absolute amounts during initial phases of decomposition.
Decomposition of SOM usually results in net releases of nitrogen, since
C : N of SOM is generally smaller than 20, much closer to C : N of microbes
than litter (Paul and Clark 1996). Degradation of proteins and nucleic acids
in SOM releases nitrogen in a mineral form (i.e., NH+4 ). The mineralized
nitrogen from SOM is partly immobilized for growth of microorganisms and
partly added to the mineral nitrogen pool in soil.
Due to the coupled carbon and nitrogen mineralization during microbial
decomposition of litter and SOM, the rate of nitrogen mineralization often
correlates with microbial respiration. For example, Zak et al. (1993) studied
carbon and nitrogen releases from labile organic matter within the forest floor
and mineral soil of Jack pine, red pine, balsam fir, sugar maple, and quaking
aspen forests in Michigan. Carbon released from microbial decomposition
was correlated with mineralized nitrogen (Nmin) by Rm = 15.9 Nmin+ 27.4 with
r = 0.853 and n = 154 for litter and Rm = 7.1 Nmin + 159.9 with r = 0.616 and
n = 154 for SOM from a laboratory incubation. Similar relationships between
net carbon and nitrogen mineralization were found in organic substrates with
low C : N ratios (Gilmore et al. 1985, Moorhead et al. 1987, Ruess and Seagle
1994, Eriksen and Jensen 2001). Across different types of soils from three
communities in an Alaskan boreal forest, rates of soil respiration were associ-
ated with rates of microbial turnover and nitrogen mineralization in a labora-
tory incubation study (Vance and Chapin 2001). In the field research, nitrogen
mineralization may not be well correlated with soil respiration due to the
nitrogen immobilization.
2.3. SOIL RESPIRATION AND REGIONAL AND GLOBAL

CARBON CYCLING
Soil respiration plays a critical role in the regulation of carbon cycling on

regional and global scales. The carbon cycle on the global scale involves
exchanges of CO2 among the land biosphere, the atmosphere, oceans, and the
earth’s crust (Fig. 2.3). Each year, photosynthesis of land plants takes up
approximately 120 Pg (1015 g) C yr−1 from the atmosphere. A similar amount of
carbon is released back to the atmosphere through ecosystem respiration.
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Meinhold, Die Bernsteinhexe. 592-594.
Mendelssohn, Phädon. 528. 529.
Möser, Patriotische Phantasien. 422-424.
Müllner, Die Schuld. 595. 596.
Münchhausens Reisen u. Abenteuer. 300. 301.
Musäus, Volksmärchen I. 225. 226.
– Volksmärchen II. 227. 228.
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Neugriechische Gedichte. 619.
Novalis, Heinrich von Ofterdingen. 497. 498.
Oehlenschläger, Corregio. 469. 470.
Pestalozzi, Lienhard und Gertrud. 315-320.
Petöfi, Gedichte. 645-647.
Platen, Die Abbassiden. 630. 631.
– Gedichte. 269. 270.
Puschkin, Boris Godunof. 293.
Racine, Britannicus. 409.
– Phädra. 440.
Raimund, Der Bauer als Millionär. 436.
– Der Verschwender. 437. 438.
Raupach, Der Müller und sein Kind. 435.
Römische Lyriker, Ausgewählte Gedichte. 578. 579.
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Sallet, Laien-Evangelium. 487-490.
– Schön Irla. 511.
Schenkendorf, Gedichte. 336. 337.
Schiller, Der Neffe als Onkel. 456.
– Turandot. 612. 613.
– Über naive und sentimentalische Dichtungen. 346. 347.
Schlegel, Englisches und spanisches Theater. 356-358.
– Griechisches und römisches Theater. 353-355.
Schleiermacher, Monologe. 468.
Schubart, Leben und Gesinnungen. 491-493.
Schwab, Doktor Faustus. 405.
– Fortunatus und seine Söhne. 401. 402.
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– Die schöne Melusina. 284.
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Tasso, Das befreite Jerusalem. 684-690.
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– Der tote Gast. 361. 362.
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Full Soil Respiration and The Environment 1st Edition Yiqi Luo Ebook All Chapters

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The Case for The Enlightenment Scotland and Naples 1680

Yiqi Luo and Xuhui Zhou

AMSTERDAM • BOSTON • HEIDELBERG

This book is printed on acid-free paper.

Copyright © 2006, Elsevier, Inc. All rights reserved.

Library of Congress Cataloging-in-Publication Data

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1. Introduction and Overview 3

2. Importance and Roles of Soil Respiration 17

3. Processes of CO2 Production in Soil 35

3.2. Root respiration 42

4. Processes of CO2 Transport from Soil to the Atmosphere 61

Part III Regulation 77

6. Temporal and Spatial Variations in Soil Respiration 107

Part IV Approaches 159

8. Methods of Measurements and Estimations 161

9. Separation of Source Components of Soil Respiration 187

Bomb 14C tracer 204

10. Modeling Synthesis and Analysis 215

Soil respiration is an ecosystem process that releases carbon dioxide from

Yiqi Luo and Xuhui Zhou

1.1. Deﬁnition and introduction 4

1.1. DEFINITION AND INTRODUCTION

CO2, depending on conditions that inﬂuence reaction equilibriums (see

Soil respiration (µmol m -2 s-1) 6

1.2. HISTORY OF RESEARCH

Soil metabolism Soil fertility Ecosystems No tillage Global change

1850 1900 1950 2000

respiration, and SOM dynamics. Schlesinger (1977) reviewed many studies

1.3. OVERVIEW OF THE BOOK

This book, which comprises 10 chapters, is dedicated to providing an under-

concentration along soil vertical proﬁles and determined by diffusion and

2.1. Soil respiration and ecosystem carbon

2.1. SOIL RESPIRATION AND ECOSYSTEM

Year Rs Re R s/Re GPP* R s/GPP NPP NEP

1999 1116 1538 0.73 1637 0.68 656 99

Equation 2.3 is a quantitative basis of the biometrical approach to estima-

Litterfall (g C m-2 yr-1)

Soil CO2 release 1263 1489 1293 1576

2.2. SOIL RESPIRATION AND NUTRIENT CYCLING

A major component of soil respiration is from microbial decomposition of

increase during the decomposition. The absolute amount of nitrogen increases

2.3. SOIL RESPIRATION AND REGIONAL AND GLOBAL

Soil respiration plays a critical role in the regulation of carbon cycling on

Vollständige Verzeichnisse sind durch jede Buchhandlung gratis zu beziehen. Die

Druck vom Bibliographischen Institut in Leipzig.