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Agroecosystems in a changing climate 1st Edition Edited
By Paul C . D . Newton Digital Instant Download
Author(s): Edited by Paul C . D . Newton , R . Andrew Carran , Grant R .
Edwards , and Pascal A . Niklaus
ISBN(s): 9780849320880, 0849320887
Edition: 1
File Details: PDF, 16.30 MB
Year: 2007
Language: english
 Agroecosystems
in a

Changing Climate

© 2007 by Taylor & Francis Group, LLC

Advances in Agroecology
Series Editor: Clive A. Edwards
Agroecosystem Sustainability: Developing Practical Strategies
Stephen R. Gliessman
Agroforestry in Sustainable Agricultural Systems
Louise E. Buck, James P. Lassoie, and Erick C.M. Fernandes
Biodiversity in Agroecosystems
Wanda Williams Collins and Calvin O. Qualset
Interactions Between Agroecosystems and Rural Communities
Cornelia Flora
Landscape Ecology in Agroecosystems Management
Lech Ryszkowski
Multi-Scale Integrated Analysis of Agroecosystems
Mario Giampietro
Soil Ecology in Sustainable Agricultural Systems
Lijbert Brussaard and Ronald Ferrera-Cerrato
Soil Organic Matter in Sustainable Agriculture
Fred Magdoff and Ray R. Weil
Soil Tillage in Agroecosystems
Adel El Titi
Structure and Function in Agroecosystem Design and Management
Masae Shiyomi and Hiroshi Koizumi
Tropical Agroecosystems
John H. Vandermeer

Advisory Board
Editor-in-Chief
Clive A. Edwards
The Ohio State University, Columbus, OH
Editorial Board
Miguel Altieri, University of California, Berkeley, CA
Lijbert Brussaard, Agricultural University, Wageningen, The Netherlands
David Coleman, University of Georgia, Athens, GA
D.A. Crossley, Jr., University of Georgia, Athens, GA
Adel El-Titi, Stuttgart, Germany
Charles A. Francis, University of Nebraska, Lincoln, NE
Stephen R. Gliessman, University of California, Santa Cruz, CA
Thurman Grove, North Carolina State University, Raleigh, NC
Maurizio Paoletti, University of Padova, Padova, Italy
David Pimentel, Cornell University, Ithaca, NY
Masae Shiyomi, Ibaraki University, Mito, Japan
Sir Colin R.W. Spedding, Berkshire, England
Moham K. Wali, The Ohio State University, Columbus, OH

© 2007 by Taylor & Francis Group, LLC

 Agroecosystems
in a

Changing Climate

Edited by

Paul C. D. Newton
R. Andrew Carran
Grant R. Edwards
Pascal A. Niklaus

© 2007 by Taylor & Francis Group, LLC

CRC Press
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Boca Raton, FL 33487‑2742
© 2007 by Taylor & Francis Group, LLC
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No claim to original U.S. Government works

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Library of Congress Cataloging‑in‑Publication Data

Agroecosystems in a changing climate / editors: Paul C.D. Newman ... [et al.].
p. cm. ‑‑ (Advances in agroecology)
Includes bibliographical references and index.
ISBN‑13: 978‑0‑8493‑2088‑0 (alk. paper)
1. Agricultural ecology. 2. Climatic changes. 3. Agriculture‑‑Environmental
aspects. I. Newman, Paul C. D. II. Series.

S589.7.A479 2006
577.5’522‑‑dc22 2006010178

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2088_C000.fm Page v Tuesday, July 25, 2006 1:15 PM

Preface
This book is the 12th in the continuing CRC Series on Advances in Agroecology.
While other volumes have discussed aspects of climate change, this is the first to
deal directly with this topic. In this book we employ a broader definition of climate
change to include changes not only in climatic factors per se (temperature and
rainfall) but also in the composition of the atmosphere (carbon dioxide in particular
but also ozone).
Climate change is an issue that engages many more participants than just the
scientific research community. The issue is highly politicised and widely presented
and discussed in a range of media and fora. It is therefore not surprising that the
opinions people hold about climate change are informed by a range of material of
which original scientific research might be only a small component. We are intro-
ducing this book into this lively arena because as experimental scientists our expe-
rience demonstrates to us that changes in temperature, precipitation, and atmospheric
carbon dioxide have the potential to profoundly alter terrestrial ecosystems and the
delivery of the services they provide. We do not have to wait for accurate projections
of a future climate to make progress here. An important task is to develop our
understanding of the effects of the climate change drivers and their interactions on
biological systems; from this base of knowledge we will be much better placed to
consider the range of future environments that may arise and the range of agroeco-
systems we will need to cover. Consequently, we have organised the book so that a
fundamental understanding of processes is presented; we have then asked applied
scientists to consider the consequences of a change in these processes for agroeco-
systems.
This book has taken a long while to prepare — a period sufficiently long for
atmospheric CO2 to increase by 6 ppm — but this event has allowed our authors to
include the most recent findings and views and we would like to thank them for
their patience and for sharing their ideas as well as their knowledge of their particular
subject areas. We would like to thank the editor of the “Agroecology” series, Clive
Edwards, and Taylor & Francis editor, John Sulzycki, for their invitation to prepare
this book and for their advice and encouragement. Pat Roberson and Linda Manis
have provided the essential publishing expertise and a number of colleagues gave up
their time to referee chapters and we thank them all for these essential contributions.

Paul C.D. Newton, R. Andrew Carran,

Grant R. Edwards, and Pascal A. Niklaus

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2088_C000.fm Page vii Tuesday, July 25, 2006 1:15 PM

Editors
Paul C.D. Newton, Ph.D., is senior research scientist in the Land and Environmental
Management group of AgResearch based in Palmerston North, New Zealand. After
completing a diploma in communications studies at the Central London Polytechnic
he worked in a number of areas including agriculture and entertainment. He then
earned a degree in agricultural botany at the University College of North Wales,
Bangor and subsequently earned a Ph.D. in a joint project between Bangor and the
Weed Research Organisation. Dr. Newton moved to New Zealand in 1986 for a
postdoctoral fellowship and has remained there, studying the effects of global change
on grazed pastoral systems. His research has included locating and working at natural
CO2 springs and the construction of a Free Air Carbon Dioxide Enrichment (FACE)
experiment in 1997. This experiment is continuing and his work concentrates on
how the relationships between plant community structure and ecosystem function
is modified by elevated atmospheric CO2.

R. Andrew Carran, is a senior research scientist at AgResearch located in Palm-

erston North, New Zealand. After graduating with a M. Agric. Sci (Hons soil science)
he worked at research stations of the Grasslands Division of the New Zealand
Department of Scientific and Industrial Research until 1991. During this period, he
researched soil processes including nitrogen fixation, acidification, and ammonia
volatilisation in sheep production systems. Through 1990, he held a Visiting
Researcher position at the Leopold Center for Sustainable Agriculture at Iowa State
University. Since 1991 his research programme has expanded to include source sink
studies of nitrous oxide and methane in soils, and the impacts of elevated CO2 on
soil processes influencing nutrient availability and loss in grazed pastures.

Grant R. Edwards, Ph.D., is a senior lecturer in Pasture Ecology in the Agriculture

and Life Sciences Division, Lincoln University, New Zealand. He completed his
undergraduate degree in Agriculture Science at Lincoln University, before being
awarded a Rhodes Scholarship to study at Oxford University. He was awarded a
DPhil from Oxford University in 1994 for his thesis titled: The Creation and Main-
tenance of Spatial Heterogeneity in Grasslands: The Role of Plant-Herbivore Inter-
actions. Thereafter, he had two postdoctoral research positions at Imperial College,
London and at AgResearch, Hamilton, New Zealand. From 2000 to 2004, he was a
lecturer at Imperial College, London. In 2004 he obtained his present position at
Lincoln University. His research interests include the ecology and management of
temperate grasslands under a changing climate. Current research conducted by his
group addresses how an understanding of livestock diet selection and plant popula-
tion dynamics can be used to manipulate the botanical composition of pastures, be
it for weed control or livestock production reasons.

© 2007 by Taylor & Francis Group, LLC

2088_C000.fm Page viii Tuesday, July 25, 2006 1:15 PM

Pascal A. Niklaus, Ph.D., is currently a senior scientist at the Institute of Plant

Sciences of the Swiss Federal Institute of Technology in Zürich, Switzerland. He
studied physical, inorganic, and organic chemistry at the University of Basel where
he obtained his diploma in chemistry in 1992; investigating model systems for
oxygenating enzymes. He obtained his Ph.D. in botany in 1997 at the Institute of
Botany of the University of Basel with investigations of global change-effects on
grassland ecosystems. During his postdoctoral years, he also was visiting scientist
at the Institute of Arable Crops Research in Rothamsted, United Kingdom and at
Landcare Research, New Zealand where he investigated soil trace gas relations.
A main focus in Pascal Niklaus’ research is on the consequences of environ-
mental change for plant communities and the biogeochemical cycling of nutrients
and water in ecosystems; special emphasis is on interactions with soils and effects
on the ecosystem’s greenhouse gas balance (CO2, CH4, N2O).

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2088_C000.fm Page ix Tuesday, July 25, 2006 1:15 PM

Contributors
Vincent Allard Grant R. Edwards
INRA-Agronomie Agriculture Group
Fonctionnement et Gestion de Agriculure and Life Sciences
l’Ecosystème Prairial Division
France Lincoln University
Canterbury, New Zealand
Daniel J. Archambault
Laurentian University Jürg Fuhrer
Sudbury, Ontario, Canada Air Pollution/Climate Group
Agroscope FAL Reckenholz
Andrew Ash Swiss Federal Research Station for
CSIRO Sustainable Ecosystems Agroecology and Agriculture
St. Lucia, Queensland, Australia Zürich, Switzerland

Joseph C. Blankinship Oula Ghannoum

Department of Biological Sciences Centre for Plant and Food Science
Northern Arizona University University of Western Sydney
Flagstaff, Arizona Penrith South, Australia
R. Andrew Carran
Stephen L. Goldson
Land and Environmental Management
AgResearch
Group
Lincoln, New Zealand
AgResearch
Palmerston North, New Zealand
Jo E. Hossell
Sukumar Chakraborty ADAS Gleadthorpe
CSIRO Plant Industry Meden Vale, Mansfield,
Queensland Bioscience Precinct Nottingham
St. Lucia, Queensland, Australia United Kingdom

Jann P. Conroy Mark J. Hovenden

Centre for Plant and Food Science School of Plant Science
University of Western Sydney University of Tasmania
Penrith South, Australia Hobart, Tasmania, Australia

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2088_C000.fm Page x Tuesday, July 25, 2006 1:15 PM

Bruce A. Hungate William H. Schlesinger

Department of Biological Sciences and Nicholas School of the Environment
Merriam-Powell Center for and Earth Sciences
Environmental Research Duke University
Northern Arizona University Durham, North Carolina
Flagstaff, Arizona
Matthew J. Searson
Jonathan A. Newman Australian Greenhouse Office
Department of Environmental Biology Department of the Environment and
University of Guelph Heritage
Guelph, Ontario, Canada Australia

Paul C.D. Newton Chris Stokes

Land and Environmental Management CSIRO Sustainable Ecosystems and
Group CRC for Tropical Savannas
AgResearch Management
Palmerston North, New Zealand Aitkenvale, Australia

Pascal A. Niklaus Richard B. Thomas

Institute of Plant Sciences Department of Biology
ETH Zürich West Virginia University
Zürich, Switzerland Morgantown, West Virginia

Matthias C. Rillig Skip J. Van Bloem

Microbial Ecology Program Depto Agronomîa y Suelos
Division of Biological Sciences Universidad de Puerto Rico
The University of Montana Mayagüez, Puerto Rico
Missoula, Montana
Lewis H. Ziska
G. Brett Runion Crop Systems and Global Change
Soil Dynamics Laboratory Laboratory
USDA-ARS USDA-ARS
Auburn, Alabama Beltsville, Maryland

© 2007 by Taylor & Francis Group, LLC

2088_C000.fm Page xi Tuesday, July 25, 2006 1:15 PM

Table of Contents
Chapter 1
Introduction................................................................................................................1
Paul C.D. Newton, R. Andrew Carran,
Grant R. Edwards, and Pascal A. Niklaus

PART I Resource Supply and Demand

Chapter 2
Climate Change Effects on Biogeochemical Cycles, Nutrients, and
Water Supply ...........................................................................................................11
Pascal A. Niklaus

Chapter 3
Nutrient and Water Demands of Plants under Global Climate Change.................53
Oula Ghannoum, Matthew J. Searson, and Jann P. Conroy

Chapter 4
Climate Change and Symbiotic Nitrogen Fixation in Agroecosystems.................85
Richard B. Thomas, Skip J. Van Bloem, and William H. Schlesinger

Chapter 5
Belowground Food Webs in a Changing Climate ................................................117
Joseph C. Blankinship and Bruce A. Hungate

Chapter 6
Herbivory and Nutrient Cycling............................................................................151
R. Andrew Carran and Vincent Allard

Chapter 7
Sustainability of Crop Production Systems under Climate Change.....................167
Jürg Fuhrer

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PART II Pests, Weeds, and Diseases

Chapter 8
Plant Performance and Implications for Plant Population
Dynamics and Species Composition in a Changing Climate...............................189
Grant R. Edwards and Paul C.D. Newton

Chapter 9
Climate Change Effects on Fungi in Agroecosystems .........................................211
Matthias C. Rillig

Chapter 10
Trophic Interactions and Climate Change ............................................................231
Jonathan A. Newman

Chapter 11
Future Weed, Pest, and Disease Problems for Plants ...........................................261
Lewis H. Ziska and G. Brett Runion

PART III Capacity to Adapt

Chapter 12
Distinguishing between Acclimation and Adaptation...........................................291
Mark J. Hovenden

Chapter 13
Plant Breeding for a Changing Environment........................................................309
Paul C.D. Newton and Grant R. Edwards

© 2007 by Taylor & Francis Group, LLC

2088_C000.fm Page xiii Tuesday, July 25, 2006 1:15 PM

PART IV Special Examples

In Parts I–III the authors have set out general principles determining agroecosystem
responses to global change and the consequences of these have been explored. In
this Part we present five Special Examples that bring the focus down to explore the
impacts of global change at the agroecosystem, technology, population, and regional
level.

Part I Resource Supply and Demand

Special Example 1
Impacts of Climate Change on Marginal Tropical Animal
Production Systems ...............................................................................................323
Chris Stokes and Andrew Ash

Part II Pests, Weeds, and Diseases

Special Example 2
Climate Change and Biological Control ...............................................................329
Stephen L. Goldson
Special Example 3
Efficacy of Herbicides under Elevated Temperature and CO2 .............................333
Daniel J. Archambault

Part III Capacity to Adapt

Special Example 4
Evolution of Pathogens under Elevated CO2 ........................................................337
Sukumar Chakraborty
Special Example 5
Adapting United Kingdom Agriculture to Climate Change .................................341
Jo E. Hossell

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2088_C001.fm Page 1 Monday, July 17, 2006 7:46 AM

1 Introduction
Paul C.D. Newton, R. Andrew Carran,
Grant R. Edwards, and Pascal A. Niklaus

CONTENTS

1.1 The Context......................................................................................................1

1.2 The Environmental Changes............................................................................1
1.2.1 Atmospheric CO2 Concentration .........................................................2
1.2.2 Temperature and Precipitation .............................................................4
1.3 The Structure of This Book .............................................................................5
References ................................................................................................................6

1.1 THE CONTEXT

Agricultural systems (agroecosystems) are enormously diverse in their biological
structure, in the climatic and socioeconomic drivers that shape them, and in the
services they provide. Agroecosystems may be critical in sustaining social, eco-
nomic, and cultural fabrics as well as playing a key role in community health. The
challenge we face is to maintain these services and roles during a period of rapid
environmental change — perhaps producing combinations of environmental condi-
tions that modern agriculture has not previously experienced. In the first instance,
this is an issue for biologists; it is essential to improve our understanding of the
potential impacts of environmental change. However, it is also the case that agro-
ecosystems are the site of intense interaction between humans and the natural world.
In some situations this offers the prospect of effective adaptation to climatic change
— either amelioration of negative effects or exploitation of positive effects; however,
at the other extreme there will be many situations in which the effects of climate
change will be exacerbated by socioeconomic factors such as income inequality or
credit availability (Patt et al. 2005). As biologists we cannot ignore this reality, and
if we want our work to be relevant, then we need to consider carefully who and how
we are targeting with our knowledge. We consider some of these issues later in this
chapter, but first we review the main drivers of environmental change.

1.2 THE ENVIRONMENTAL CHANGES

The changes in the environment we are most concerned with involve temperature,
precipitation, and atmospheric CO2 concentration. Here we consider some of the

© 2007 by Taylor & Francis Group, LLC

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2 Agroecosystems in a Changing Climate

issues surrounding these drivers of change, which we feel are of central importance
in determining biological responses. In particular we draw attention to variation in
current and predicted temperature and precipitation trends, and to the likelihood of
further increases in atmospheric CO2 concentration.

1.2.1 ATMOSPHERIC CO2 CONCENTRATION

As well as its indirect effect on ecosystems through its role as a greenhouse gas,
atmospheric CO2 has a direct effect on the biosphere because it is the primary raw
material for plant growth. Direct measurements of atmospheric CO2 have been made
since 1958 in Hawaii (Keeling et al. 1982), and new monitoring stations have been
established since that time to give a more global coverage. The data from these
stations (CDIAC 2005), together with historical records extracted from ice cores
(CDIAC 2005), show the atmospheric concentration has increased exponentially
since the late 19th century, that it increases each year, and that it is now at a level
higher than at any time in the last 650,000 years (Siegenthaler et al. 2005).
Because CO2 is such an important driver of biological systems, our picture of
the future must include the rate of change in atmospheric CO2 and the level at which
the concentration will stabilize; these both depend upon the balance between CO2
emissions and the rate at which CO2 can be sequestered into sinks. A recent carbon
(C) budget for the 1990s (Schimel et al. 2001) calculates emissions from fossil fuels
to be 6.3 Gt C yr –1, with an additional 1.7 Gt C added from land use change.
Approximately half of CO2 emissions are removed by oceanic and terrestrial sinks
leaving an annual addition of approximately 3.2 Gt C to the atmosphere. During
this period, the CO2 concentration in the atmosphere increased at a rate of about 1.5
ppm or 0.44% per year. If this balance of sources and sinks were to continue, then
by 2050 the concentration in the atmosphere would reach 460 ppm. As neither sink
nor source activity is expected to stay constant over this period, prediction becomes
a difficult task.
On the source side, prediction of future emissions depends upon a range of
assumptions about technology change and population growth. Emissions projected
for 2050 range from 11.0 to 23.9 Gt C (Prentice et al. 2001). To put emissions
reduction in perspective, we can make a crude calculation: Assuming current emis-
sions of 6.3 Gt yr –1 (Schimel et al. 2001) and a world population of 6 billion, then
our current emissions rate is 1 t of C per capita yr –1. To maintain a balance with
the sinks, which currently absorb only half of these emissions, and to have no net
increase in the atmosphere, we need to have emissions of 500 kg per capita yr –1. If
we look at current rates of per capita emissions of C from fossil fuel use (Marland
et al. 2004), we find that emissions from the United States and Western Europe are,
respectively, 10 times and 4 times the 500 kg rate that would balance current sinks.
The current per capita emission rate in China also exceeds the “stabilising” output
level, and it is only in countries such as Africa and India that emissions are below
the 500 kg per capita level. These figures suggest that major changes in our energy
creation and use will be necessary to achieve significant reductions in emissions
(Hoffert et al. 2002).

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Introduction 3

The second half of the equation is the activity and size of sinks. Again, we can
make a rough calculation to establish the size of the problem by considering how
much sink activity would need to change by 2050 to keep net emissions to the
atmosphere at the current 3.2 Gt C level; that is, to constrain the annual increase to
1.5 ppm. Using the low prediction of CO2 emissions for 2050 of 11 Gt C (Prentice
et al. 2001) would require sinks to remove 7.8 Gt C to maintain net emissions at
3.2 Gt C. If half of the sink activity is oceanic and half terrestrial, then terrestrial
sinks would need to absorb 3.9 Gt C or 2.8 times their current rate (assuming a
current terrestrial sink of 1.4 Gt yr –1, Prentice et al. 2001). The stimulation of plant
growth due to the rising concentration of CO2 offers the promise of enhanced sink
strength; however, in the absence of significant changes in the ratio of C to nitrogen
(N) in terrestrial pools, such an increase in C sequestration would require substantial
increases in N availability, perhaps beyond the capacity of ecosystems to provide
(Hungate et al. 2003). In fact, elevated CO2 may exacerbate this constraint, as a
common response appears to be a progressive decline in the availability of N to
plants (Luo et al. 2004). The potential sink capacity of the terrestrial biosphere
remains a critical value if we are to predict future CO2 concentrations. However, a
doubling of sink capacity would be required to absorb even current emissions, let
alone those expected in the next decades, placing an unrealistic expectation on the
absorbing capacity of this sink.
The average annual CO2 concentration of the well-mixed atmosphere does not
differ greatly among monitoring stations, although there is a slightly lower average
in the Southern Hemisphere. However, at different scales there can be considerable
variation in concentration, and it is relevant to consider whether these variations are
likely to change in the future in response to the changing climate and atmosphere.
The net CO2 exchange of the biosphere results in marked seasonal differences (15
to 20 ppm, Keeling et al. 1996) in atmospheric concentration in the Northern
Hemisphere; interestingly, the amplitude of this difference is increasing over time
(Keeling et al. 1996), probably because of disturbance and a change in the identity
and activity of the vegetation (Zimov et al. 1999). Regional differences in atmo-
spheric CO2 concentration can arise from urban development where large sources
of fossil fuel use can dominate the concentration profile. For example, Ziska et al.
(2004) measured average concentrations of 466 at 0.5 km from the city centre of
Baltimore, Maryland — 401 ppm 10 km from the centre and 385 ppm at a distance
of 50 km. Temperature gradients are also established by urbanization, and clearly
both CO2 and temperature gradients will be determined by future urban development
and energy use.
Plants also experience large differences in CO2 concentration between day and
night (often > 100 ppm; e.g., Ziska et al. 2001); we are not aware of data considering
trends in this difference, but certainly one aspect of climate change has been a
reduction in the diurnal temperature range (Prentice et al. 2001), and it may be that
this could influence the biological processes of C fixation and respiration that largely
govern the differences in CO2 concentration near the surface. These biological
processes also result in considerable spatial variation in concentrations of CO2 within
plant canopies; plants growing close to the soil surface experiencing concentrations
of CO2 perhaps 100 ppm greater than plants with foliage higher in the canopy

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4 Agroecosystems in a Changing Climate

(Bazzaz and Williams 1991). It is not certain whether this spatial variation will alter
under climate change, but as soil respiration is sensitive to both temperature and
elevated CO2 (e.g., King et al. 2001) there is a strong likelihood of different canopy
profiles in the future. The spatial and temporal variation occurs at scales relevant to
plant growth and has been shown to influence plant responses to elevated CO2 (Ziska
et al. 2001).

1.2.2 TEMPERATURE AND PRECIPITATION

The global mean near-surface temperature record shows an increase over the 20th
century of about 0.6ºC (Folland et al. 2001), which is consistent with satellite data
for tropospheric temperatures (Tett et al. 1999; Vinnikov and Grody 2003). Further
evidence for a temperature change are the “fingerprints” of increasing temperature
that can be seen in a range of biological data such as phenological records (Parmesan
and Yohe 2003; Root et al. 2003). There are a number of forcing agents that can
modify climate, some of which are natural (solar radiation and volcanic aerosols)
and some anthropogenic, including greenhouse gases, tropospheric aerosols, cloud
changes, and changes in the land surface characteristics altering albedo (Hansen et
al. 1998). When climate models are run to simulate long-term temperature trends,
the prediction for the latter part of the last century requires greenhouse gas effects
be included in order to adequately simulate the observed changes (e.g., Karoly et
al. 2003); studies such as these are part of the argument that anthropogenic emissions
of greenhouse gases are resulting in a change in our climate (Mitchell et al. 2001).
Predictions for the change in mean temperature over the next 100 years range
between 1.4 and 5.8ºC (Cubasch et al. 2001). Spatial variation in temperature and
precipitation trends are widely observed and predicted. Rainfall has increased by 10
to 40% over the past 100 years in northern Europe, but has decreased by 20% in
southern Europe. In the United States, soil temperatures (1 m over the period 1967
through 2002) show a positive warming trend at stations in the north and northwest-
ern United States, but a strong cooling trend in the southeastern part of the country.
Because part of the spatial variation is driven by land use (Pielke et al. 2002; Stone
and Weaver 2003; Feddema et al. 2005), variation within regions is also apparent.
For example, Pielke et al. (2002) examined the long-term records from a cluster of
stations in eastern Colorado and found “enormous” differences, defying attempts to
calculate regional trends.
Any trend in mean annual temperature or precipitation is unlikely to be evenly
distributed across seasons. For example, in Australia there has been a trend since
1950 for an increase in minimum temperatures in all seasons in Queensland, but a
cooling of maximum summer temperature in northwestern Australia (Anonymous
2005). Differences can be a matter of degree, such as the long-term increase in
European summer temperatures over the past 100 years of 0.7ºC compared to an
increase of 1.1ºC in winter temperatures; or can be quite strikingly different even
in sign, such as the trends in soil temperatures (40 cm) at Irkutsk, Russia, where
there was a marked positive trend in annual average temperature over the past 120
years, but a decline in the average summer temperature of 4ºC over this time (Zhang
et al. (2001).

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Introduction 5

Variation is also evident at shorter timescales with a reduction in the diurnal

temperature range being frequently and widely observed (Prentice et al. 2001). As
one of the major factors implicated in this change is soil moisture content (Stone
and Weaver 2003), there is a direct link back to vegetation responses to the changing
environment. Diurnal patterns in precipitation have been less thoroughly investi-
gated, but here again trends have been identified (Dai 1999).

1.3 THE STRUCTURE OF THIS BOOK

In summary, we can anticipate a continuing increase in the global mean atmospheric
CO2 concentration, and in global mean temperature with a variety of changes in
temperature, precipitation, and CO2 occurring at different scales relevant to biolog-
ical activity. How can we deal with this complexity?
We suggest that two aspects are particularly important. First, an improved under-
standing of the biological consequences achieved through greater integration of basic
and applied knowledge; and second, a clearer focus on the audience for this research,
as this should enable us to ask and address more targeted and relevant questions.
Our perception is that climate change impact research often occurs in two ways.
On the one hand there are studies of the direct effects of a particular set of climate
drivers on a specific crop, often concentrating on the agricultural outputs, such as
yield and quality (e.g., Reddy and Hodges 2000). On the other hand, there is a more
ecological literature that seeks to find some general principles of response (e.g.,
Körner and Bazzaz 1996, p. 4). In this volume we hope to draw these two approaches
together so that ecologists can provide the “theoretical underpinning that informs
them (agriculturalists) what might be happening, what to look for, and what to build
on” (Lawton 1996, p. 4), and agriculturalists can interpret these ecological insights
and general theory in relation to agroecosystem performance. Consequently, each
section of this book combines general principles of response leading to applied
consequences. We have sections considering (1) the supply of resources necessary
to sustain agriculture in the future, which we identify from an understanding of how
climate change will modify biogeochemical cycles and changes in plant nutrient
demands; (2) the incidence of pests, weeds, and diseases and their control for which
we need an understanding of how the population biology of organisms will change;
and (3) the adaptations that might be possible, including plant breeding solutions,
for which we need an understanding of the capacity for adaptation that exists in
plant populations. In addition to the full chapters, we have included Special Example
chapters that deal in more detail with specific issues.
Having collected the best information, we are then faced with the issue of
communicating it effectively to interested groups and, in particular, to those groups
that can act effectively in leading or implementing adaptive measures. On this basis
we suggest that while farmers and landowners will likely be interested in projections
that consider, for example, changes in yield or the incidence of pests, their behaviour
is unlikely to be modified by such predictions, as they tend to be responsive to
current conditions. Consequently, we imagine that the issues in this book will
resonate most strongly with other researchers and with agribusiness because here
are important messages about potential opportunities for the development of new

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6 Agroecosystems in a Changing Climate

technologies. These are important sectors to reach, as it is through the development

of new, adaptive technologies that we can imagine making a difference in agroeco-
system performance in a changing environment.

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8 Agroecosystems in a Changing Climate

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Part I
Resource Supply and Demand

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2 Climate Change Effects

on Biogeochemical
Cycles, Nutrients, and
Water Supply
Pascal A. Niklaus

CONTENTS

2.1 Introduction ....................................................................................................12

2.1.1 Essential Elements .............................................................................12
2.1.2 Mechanisms and Key Processes ........................................................14
2.1.2.1 Nutrient Balance of the Whole Ecosystem ........................15
2.1.2.2 Mineralisation of Soil Organic Matter ...............................16
2.1.2.3 Immobilisation in Soil Microbial Biomass ........................16
2.1.2.4 Leaching and Volatilisation of Nutrients............................17
2.2 Effects of Elevated CO2 on Nutrient Cycling ...............................................17
2.2.1 Evidence for Changes in Soil Carbon Fluxes ...................................17
2.2.2 Nutrient Immobilisation and Mineralisation
Responses to Elevated CO2................................................................18
2.2.3 Plant Tissue Quality ...........................................................................22
2.2.4 Leaching and Volatilisation................................................................27
2.3 Effects of Elevated Temperature and Precipitation on
Nutrient Cycling.............................................................................................28
2.3.1 Net Primary Production, Decomposition, and the
Carbon and Nitrogen Balance of Ecosystems...................................30
2.3.2 Nutrient Immobilisation and Mineralisation .....................................33
2.3.3 Plant Tissue and Litter Quality..........................................................35
2.3.4 Leaching and Erosion ........................................................................36
2.4 The Hydrological Cycle and Scaling Issues .................................................37
2.5 Conclusions ....................................................................................................40
2.5.1 Multiple Element Interactions ...........................................................40
2.5.2 Soil Types...........................................................................................41
2.5.3 Controls of Crops and Forage Nutrient Contents .............................41
2.5.4 Integrated Studies along Food Chains...............................................41
2.5.5 Multiyear Ecosystem-Level Studies ..................................................42

11

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12 Agroecosystems in a Changing Climate

2.5.6 Hydrological Feedback ......................................................................42

Acknowledgments....................................................................................................42
References................................................................................................................42

2.1 INTRODUCTION
The atmospheric CO2 concentration has increased by ≈30% relative to the preindustrial
concentration of 280 µL L–1 and is projected to reach 540 to 970 µL L–1 by the end of
this century, depending on emission scenarios and climate feedback (IPCC [Intergov-
ernmental Panel on Climate Change], 2001). CO2 and other atmospheric gases of
anthropogenic origin are radiatively active, and increases in global temperatures in the
range of 1.4 to 5.8° C are predicted, depending on emission scenarios and climate
sensitivity (IPCC, 2001). As a consequence of warming, changes in the global distri-
bution of precipitation are anticipated, with projected increases at medium to high
latitudes, but decreases in other areas (e.g., the European Mediterranean).
In this chapter, I analyse how these global changes might affect the biogeochem-
ical cycling of nutrients and hydrology, and how this ultimately may impact on
agricultural ecosystems.
An important distinction to be made is between agroecosystems with high
fertiliser input and relatively open nutrient cycles on one hand, and low input and
seminatural systems in which nutrient cycles are relatively closed on the other hand.
In intensified agriculture, relatively large amounts of nutrients are removed from the
ecosystem with the crop and need to be resupplied in the form of mineral or organic
fertiliser. Effects of global change on soil nutrient cycling are less likely to be of
importance in these systems. However, soil processes such as trace gas emissions
may change, which can strongly feed back on the climate system. Also, the nutri-
tional composition of crops may change, altering their nutritional quality and pos-
sibly requiring changes in fertiliser composition. In natural ecosystems as well as
in extensively managed systems, such as low-input crop cultivation, pastures, range-
lands, and low-intensity forestry, a significant fraction of plant nutrient demand is
met by the internal mineral nutrient cycles of soils, and effects of climate change
on soil processes may directly feed back on plant growth.
Despite several decades of global change research, available data on the effects
on soil nutrient cycling are surprisingly limited, especially for agroecosystems and
nutrients other than N. In many areas, we are still in the stage of pattern searching
without having a very detailed understanding of the mechanisms underlying
responses. In the attempt not to unnecessarily narrow the scope of this chapter to
what is already well known, included is data from all available sources, including
studies in natural ecosystems. Whether and to what extent these findings can be
extrapolated to typical agricultural situations remains to be explored.

2.1.1 ESSENTIAL ELEMENTS

Plant tissue is primarily composed of carbon (C), hydrogen (H), and oxygen (O).
These elements are derived from the fixation of atmospheric CO2 and from the

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Climate Change Effects 13

uptake of soil H2O, and are generally available in ample quantities. However, vir-
tually all naturally occurring elements are also found in plants, and more than 10
are essential for growth (Welch, 1995). Mineral nutrients are generally classified
into macronutrients, required by plants at relatively large concentrations (nitrogen
[N], phosphorus [P], potassium [K], sulfur [S], calcium [Ca] and magnesium [Mg];
Epstein, 1965); and micronutrients, which are required in much lower quantities
(chlorine [Cl], iron [Fe], boron [B], manganese [Mn], zinc [Zn], copper [Cu],
molybdenum [Mo] and nickel [Ni]). Still other elements are beneficial to plants but
probably not essential for growth (sodium [Na], silicon [Si], cobalt [Co] and sele-
nium [Se]). Micronutrients are predominantly bound in enzymes, where they often
have important functional roles at the active sites, whereas macronutrients are con-
stituents of organic macromolecules (e.g., N, P, and S in proteins and nucleic acids)
or act as osmotica (e.g., K).
Most studies of global change effects on nutrient cycling have so far focused
on nitrogen. One reason may be that N is the nutrient required in the largest quantity;
another reason may be the relative ease with which N can be measured. The nitrogen
cycle is also clearly the most complex of all cycles of essential elements because N
occurs at a wide range of oxidation states; is involved in a vast array of microbial
transformations; and also occurs in gaseous, solid, and dissolved forms, endowing
it with exceptional mobility. There is, therefore, a large potential for climate change
to interfere with N nutrition. However, the N cycle is also special in that a biological
pathway exists with N2 fixation by which ecosystems can adjust to altered N demand.
This is not the case for the other elements. Indeed, while N clearly is often limiting
(Vitousek and Howarth, 1991), the level at which N becomes limiting in the long
term is frequently determined by the availability of other mineral nutrients (e.g., P:
Cole and Heil, 1981; McGill and Cole, 1981; Niklaus and Körner, 2004, Mo:
Hungate et al., 2004).
Many animals, both wild and domestic, forage on plants and accommodate their
mineral nutrient needs from plant sources. Animals require many of the same
elements as plants, but additionally require various complex organic molecules. Plant
chemical composition, therefore, can determine animal growth, but the limiting
component is often not easy to determine. For example, herbivores are often more
limited by N than by carbohydrates. An important consideration is that nutrient
concentrations that are sufficient for plants may be too low for the animals that feed
on them. For example, many New Zealand and Australian soils are very low in
cobalt; cobalt is not essential to plants* and, thus, does not limit their growth.
However, sheep foraging on these plants exhibit severe cobalt-deficiency symptoms
(Lee et al., 1999). The accumulation of nonplant-essential elements in plant tissue
(e.g., iodine or cobalt) and the accumulation of plant-essential elements beyond
limiting concentrations, therefore, can be ecologically and economically important.
Human nutrition ultimately also depends on plant chemical composition,
whether plants are consumed directly or indirectly as animals that previously fed
on plants (Underwood and Mertz, 1987). Besides insufficient total energy and protein
input, the World Health Organization (WHO) identified micronutrient deficiency as

* Except for N2 fixation.

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14 Agroecosystems in a Changing Climate

a major cause of malnutrition (WHO, 2000). Over 2 billion people are currently
affected by deficiencies in iodine, zinc, iron, selenium, and calcium, but also of more
complex phytochemicals such as vitamins A and E, niacin, and folate (Grusak and
DellaPenna, 1999; WHO, 2000). Clearly, human malnutrition at the global level is
a complex phenomenon involving political, sociological, and economic aspects
beyond the scope of this chapter. However, these data emphasize that micronutrient
effects of global change may have important implications beyond the functioning
of plants (Allaway, 1987).

2.1.2 MECHANISMS AND KEY PROCESSES

In the following analysis, I focus on a number of key processes in nutrient cycles
that are experimentally accessible. I review experimental evidence on how these
respond to simulated climate change, and analyse, from a theoretical point of view,
how they may be affected, directly or indirectly, by elevated CO2, elevated temper-
atures and altered precipitation. While these drivers are quite different and the details
of the mechanisms involved clearly are complex, two principal groups of mecha-
nisms can be identified (Figure 2.1).
A first group of effects is related to changes in carbon cycling. A primary effect
of elevated CO2 is to increase photosynthesis and plant growth; warming also
increases primary production of most ecosystems, but it also affects respiration and
decomposition. Elevated CO2 and warming, therefore, result in alterations of the
carbon balance of plants, soils, and soil organisms.
The second group of effects is related to the hydrological cycle. Elevated CO2
reduces stomatal conductance in virtually all vascular plant species, and this can
reduce evapotranspiration and water use, at least per unit of plant biomass. Warming,
on the other hand, increases evapotranspiration and generally results in a drier
environment. Precipitation, finally, has a direct effect on the water balance of the

Global Change Primary Processes Net Effects on Key Processes

Drivers Affected Effects of Nutrient Cycles

FIGURE 2.1 Schematic of effects of global change drivers via alterations of carbon and water
balance. See text for a detailed discussion.

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Climate Change Effects 15

FIGURE 2.2 Key processes in nutrient cycles that might be affected by global environmental
change (SOM = soil organic matter).

ecosystem. Alterations of the ecosystem water balance affect many processes includ-
ing the diffusion of gases and nutrients, sorption processes, and leaching rates. Redox
potential is also strongly affected by the soil water balance, primarily by controlling
gas-filled pore space and, therefore, the diffusion of O2; this can induce shifts in the
activity of different microbial groups involved in nutrient transformations, but oper-
ating under different redox conditions, for example, between nitrifying and denitri-
fying bacteria.
Effects via carbon and water balance interact because in many ecosystems
productivity and decomposition are limited by low soil water content (but by high
soil moisture in water-logged soils). There are also more complex feedback mech-
anisms involved that will be discussed. Some key quantities and processes that are
focused on in this chapter are schematically shown in Figure 2.2.

2.1.2.1 Nutrient Balance of the Whole Ecosystem

Summing all nutrient fluxes across an ecosystem’s boundaries theoretically allows

calculation of the nutrient balance of the whole ecosystem. If nutrient outputs are
larger than inputs, nutrient stocks within the ecosystem will become depleted and
productivity may decrease in the long run. Data allowing for a complete balance are
scarce; however, there is a substantial body of literature reporting the main fluxes,
and these may give an indication of the changes to expect in a future climate.
Climate change may affect the nutrient balance of ecosystems in several ways:

1. Nutrient deposition from the atmosphere could change with altered

weather patterns (precipitation, range of transport, anthropogenic emis-
sions of pollutants [N, S], etc.).
2. Biological fixation of atmospheric N2 may change (addressed by Thomas
et al., Chapter 4, this volume).
3. Weathering rates of minerals may change.

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16 Agroecosystems in a Changing Climate

4. The amount of nutrients exported may change due to changes in amount

and elemental composition of plant material.
5. Losses by leaching and erosion could change due to effects on nutrient
cycling and alterations in climate, especially in precipitation.

In the long run, changes in the nutrient balance may necessitate a corresponding
change in the elemental ratio of nutrient inputs to maintain the status quo.

2.1.2.2 Mineralisation of Soil Organic Matter

Plants take up nutrients from soil solution. One process delivering nutrients to soil
solution is the mineralisation of organic matter. The mineralisation of different
elements does not occur in concert because mineral nutrients differ in the way they
are bound in organic matter (McGill and Cole, 1981). Nitrogen is predominantly
covalently bound to carbon. Sulphur is covalently bound both directly to C and via
ester linkages. Phosphorus is essentially ester-bound, while potassium does not bind
covalently at all, but forms ionic bonds. The mineralisation of N is, therefore, largely
coupled to that of C, while ester-linked P and S can be mineralised independently
of C by the hydrolytic action of soil exoenzymes.
Cycling rates of mineral nutrients are controlled by complex interactions
between plants, soil microbes, and abiotic factors (Schlesinger, 1996). These controls
differ between mineral nutrients: The cycling rate of limiting nutrients will control
plant productivity (e.g., N and P: Güsewell, 2004; Niklaus and Köorner, 2004;
Vitousek and Howarth, 1991; Vitousek et al., 1993), whereas plant productivity will
generally control the cycling of nutrients that are available in excess (e.g., sulphur).
The cycling of still other elements is predominantly geochemically controlled and
relatively independent of plant growth (e.g., chlorine).
Global change can alter organic matter mineralisation rates in several ways. For
example, the quality and quantity of organic matter produced by plants can be
affected, resulting in altered decomposition. Microbial activity and decomposition
can be affected by soil fluxes of plant-derived C (priming effect), and abiotic con-
ditions such as temperature and moisture, which are important controls of decom-
position rates, may also be changed.

2.1.2.3 Immobilisation in Soil Microbial Biomass

The soil microbial biomass is a highly dynamic organic matter pool and its nutrient
content often exceeds that of plants. Changes in amounts of mineral nutrients bound
in microbial cells can substantially alter the within-ecosystem nutrient distribution
and affect plant growth. The soil microbial community is predominantly saprophytic
and, therefore, depends on inputs of plant-derived organic material. As a conse-
quence, soil microbial community biomass may respond to alterations in soil C
fluxes under global change. Apart from C supply, soil microbial biomass is also
controlled top–down by grazing by protozoa and microfauna (Blankinship and
Hungate, Chapter 5, this volume), by the availability of nutrients such as P or N,
which can (co-)limit microbial growth, and by soil conditions that can have strong

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Climate Change Effects 17

effects on microbial populations (e.g., soil moisture, temperature, freeze–thaw and

drying–rewetting cycles, soil mineral, and aggregate structure).

2.1.2.4 Leaching and Volatilisation of Nutrients

Significant amounts of nutrients can be lost from ecosystems by leaching and, in

the case of N, by volatilisation of NH3, NO, N2O, and N2 (Barnard et al., 2005).
Leaching losses will generally be more important for compounds such as NO 3− and
K+, which are highly soluble and show low sorption to the solid phase. Leaching
rates are related to many factors that are likely to change in a future climate, for
example to soil solution concentrations of nutrients and the amount of water drained.
However, water flow often follows preferential flowpaths in the soil, and leaching
events are erratic, so that predictions are difficult.

2.2 EFFECTS OF ELEVATED CO2 ON NUTRIENT

CYCLING
Elevated CO2 concentrations often stimulate plant growth. One mechanism for this
response is that photosynthesis is stimulated by elevated CO2, at least in C3 plants.
A second mechanism is that in almost all species, stomatal closure is induced in
response to the increased availability of CO2. Lower stomatal conductance can result
in reduced evapotranspiration, which in turn can result in comparably higher soil
moisture at any given plant biomass, or to the maintenance of higher plant biomass
at any given level of soil H2O (Jackson et al., 1998; Niklaus and Körner, 2004;
Owensby et al., 1999). A recent analysis has demonstrated that this latter indirect
effect (increased water use efficiency) accounts for much of the biomass stimulation
observed in arid and semiarid but also in some mesic grasslands exposed to elevated
CO2 (Morgan et al., 2004b), and both C3 and C4 plants, therefore, generally exhibit
growth responses to elevated CO2 (Wand et al., 1999). Nutrient dynamics may be
altered by both increased C fixation and increased soil moisture.

2.2.1 EVIDENCE FOR CHANGES IN SOIL CARBON FLUXES

Higher plant productivity under elevated CO2 will ultimately increase organic matter
inputs to soils unless all the extra production is removed as harvested plant material
(hay, crop).* The pathways by which extra C enters the soil under elevated CO2,
however, still remain elusive. Only limited field data is available (see, for example,
Hungate et al., 1997b; Niklaus et al., 2001a; Ross et al., 1995), in part due to
methodological difficulties to assess plant–soil C fluxes in situ (Darrah, 1996; Hilbert
et al., 1987; Hungate et al., 1996; Lund et al., 1999; Niklaus et al., 2000). Exudation
is a major component of soil C inputs, but there are virtually no data on flux rates
and chemical composition of exudates under field conditions. There have been early
indications from laboratory studies, mostly conducted under hydroponic conditions,
that rhizodeposition per unit root length would increase under elevated CO2 (review

* Even then, soil organic matter inputs will increase due to biomass turnover between harvests.

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18 Agroecosystems in a Changing Climate

by Darrah, 1996), and this effect has been interpreted as plants being saturated with
carbohydrates and passively leaking or actively “disposing” belowground the extra
C not needed. However, this notion could not be corroborated in a number of field
systems, probably because soil matrix effects and resorption of organic compounds
reduce or eliminate the effects observed in hydroponics (Darrah, 1996; Jones and
Darrah, 1996). Field data are mostly based on observations of fine root turnover
(Arnone et al., 2000; Fitter et al., 1996; Pregitzer et al., 2000) and carbon tracer
studies (e.g., Hungate et al., 1997b; Newton et al., 1995; Niklaus et al., 2001a).
Nevertheless, even in the absence of a carbon overflow-type effect on rhizodeposition
under elevated CO2, and in the absence of increased root-to-shoot ratios, increased
plant productivity should result in increased soil C inputs. However, the ultimate
magnitude of this effect is still unknown, and so are the pathways by which extra
C enters the soil.

2.2.2 NUTRIENT IMMOBILISATION AND MINERALISATION RESPONSES

TO ELEVATED CO2

Increased soil C fluxes under elevated CO2 could lead to higher soil microbial
biomass and immobilisation of mineral nutrients. This negative feedback on plant
growth has been demonstrated in a pot CO2 experiment where microbial biomass
N increased and plant responses to elevated CO2 were negative (Diaz et al., 1993),
presumably due to increased input of high C:N compounds to soils. Extra C inputs
to soils can, however, increase microbial activity and, thus, prime the mineralisation
of organic matter. This positive feedback on plant growth has been proposed by Zak
et al. (1993). Priming effects on mineralisation and increased nutrient immobilisation
in microbial biomass are not mutually exclusive and microbial biomass and the
availability of nutrients to plants may increase concurrently (higher net immobili-
sation plus increased net mineralisation rates).
Experimental evidence of elevated CO2 effects on microbial biomass is equiv-
ocal. While some greenhouse studies reported increases in microbial biomass,
responses in field studies of grassland appear to be smaller and often even absent
(Table 2.1). Various experimental protocols were used; nitrogen mineralisation, for
example, was measured in the field or using isolated soil samples, with methods
that ranged from buried bags, aerobic or anaerobic laboratory incubations, to short-
and long-term 15N isotopic pool dilution methods. Despite all these differences, some
broad response patterns emerged.
Microbial C often does not respond to elevated CO2, even after several years of
increased plant productivity. However, microbial N and N mineralisation increase
in many studies, suggesting that microbial N responds more than C, and that this
extra microbial N originates from increased mineralisation of soil organic matter
rather than from immobilisation of soil mineral N. Two different mechanisms may
explain these observations: First, microbial N may have been primed by extra C
inputs under elevated CO2 (mechanism proposed by Zak et al., 1993); second,
increased soil moisture at elevated CO2 may have led to increased N mineralisation,
at least in (temporarily) H2O-limited ecosystems. CO2 effects may effectively be
indirect, that is, soil moisture effects in disguise. It is very difficult to disentangle

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Climate Change Effects 19

these two components of elevated CO2 unless an experimental design involves a

factorial irrigation treatment.
I have previously argued that cases in which microbial sequestration of extra N
under elevated CO2 occurs may be restricted to systems where N supply is abundant
and nutrient cycles are not in equilibrium with plant demand, that is, the cycling of
C and N are not strongly coupled (Niklaus and Körner, 1996); others arrived at
similar conclusions (Hu et al., 1999). The data that have become available in the
meanwhile, however, indicate that increased net microbial N immobilisation occurs
in many (though not all) systems exposed to elevated CO2. High N supply seems to
favour this response, probably via increased plant growth under elevated CO2 and
associated increased soil inputs of C and N from plants; however, this increase in
N is generally paralleled by an increase in N mineralisation rates, so that reductions
in plant N availability (the negative feedback mechanism identified by Diaz et al.,
1993) appear not to happen. One reason why this is not the case may be that overall
the C:N in organic matter inputs to soils does not change much under elevated CO2.
Dynamics of P and S might respond to elevated CO2 in a fashion similar to that
of N; after all, increases in soil microbial biomass will be accompanied by the
immobilisation of these nutrients as well, and the decomposition of soil organic
matter will release the mineral nutrients that it contained. However, an important
difference is that a significant portion of soil organic P and S is ester-bound and,
therefore, can be mineralised independently of C as has been discussed above.
Microbial biomass P and S can be measured with techniques analogous to the
ones used for N (e.g., chloroform fumigation-extraction; Brookes et al., 1982; Wu
et al., 1994); however, to my knowledge, absolutely no data is currently available
from elevated CO2 studies. At first approximation, microbial P and S pools could
be assumed to change in concert with microbial biomass, but this assumption can
be poor, as data for N demonstrate.
An indicator of P status of plants and soil microbes is the activity of soil
extracellular phosphatases; these enzymes are released in response to growth limi-
tations by plants, and their activity is especially high in P-depleted zones in the
immediate vicinity of plant roots (Barrett et al., 1998; Spiers and McGill, 1979).
Experimental data under elevated CO2 is quite limited. In a grassland, Ebersberger
et al. (2003) found a 32% increase in alkaline phosphatase activity in a N–P-colimited
calcareous meadow (Niklaus and Körner, 2004) exposed to elevated CO2 for 6 years
(but no increase in enzymes related to the N cycle). Moorhead and Linkins (1997)
reported increased phosphatase activities in tussock tundra exposed to elevated CO2
for 3 years, and Dhillion et al. (1996) found increased acid phosphatase activity in
soil turves with the annual Mediterranean grass Bromus madritensis (but no change
in extractable soil inorganic P and N). For trees and forest ecosystems, increases in
acid phosphatase activity have been found in soil below Quercus ilex exposed to
twice-ambient CO2 for 5 years (Moscatelli et al., 2001), but no effects were found
in a plantation of 16-year-old Pinus taeda exposed to elevated CO2 for 2 to 3 years.
Matamala and Schlesinger (2000) and de Lucia et al. (1997) reported signs of
decreased phosphatase activity under ponderosa pine seedlings exposed to elevated
CO2 (but increased concentrations of P-chelating oxalate).

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TABLE 2.1
Effects of Elevated CO2 on Microbial Biomass C and N, N-Mineralisation and Nitrification
Effects of CO 2 Enrichment
CO 2 Years for
Treatment Which Effects Microbial Biomass N-
System/Species Conc. (ppmv) Are Listed Fertiliser Inputs C N Mineralisation Nitrification Reference

Grassland
Lolium perenne 600 2 14 and 56 g N m–2a–1 n.s. n.s. n.s.1 Schortemeyer et al. 1996
Trifolium repens 600 2 14 and 56 g N m–2a–1 n.s. n.s. n.s.1 Schortemeyer et al. 1996
Lolium perenne 600 1 to 10 14 g N m–2a–1 n.s.2 Schneider et al. 2004
600 1 to 10 56 g N m–2a–1 ↑2 Daepp et al. 2000

Annual grassland A+350 2 to 5 unfertilised ↑/= ↑3 Hungate 1997a,b, 2000

(sandstone soil)

Agroecosystems in a Changing Climate

Annual grassland A+350 2 to 5 unfertilised ↑/= ↑3
(serpentine soil)

Perennial pasture 475 1 to 5 0 kg N m–2a–1; n.s. ↑/= ↑)

(↑ n.s. Ross et al. 2004;
P,K,Mg,S supplied pers. comm.

Perennial meadow 600 6 unfertilised n.s. ↑/= ↑/= n.s.5 Niklaus 1998; Niklaus
et al. 2001d, 2003
Planted perennial 600 5 disturbed,unfertilised n.s. n.s. = ↓5 Niklaus et al. 2001b,c
communities

Alpine meadow 680 4 unfertilised n.s. n.s. Niklaus and Körner 1996;
Körner et al. 1997
Alpine meadow 680 4 full fertiliser (equiv. n.s. n.s.
4.5 g N m–2a–1)
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2×A ↑/=4 ↑/=4

Climate Change Effects

Tallgrass prairie 2 to 8 unfertilised Williams et al. 2000;
Rice et al. 1994

Arid desert 550 3 to 5 unfertilised n.s.6 n.s. Billings et al. 2004

Shortgrass steppe 720 unfertilised n.s. n.s. Stahl, pers. comm.;

Morgan et al. 2004a

Lolium perenne/ 600 3 full fertiliser (equiv. n.s. n.s.5 Barnard et al. 2004
Holcus lanatus 24 g N m–2a–1)
Wood y
Pine forest A+200 1 to 5 unfertilised n.s. n.s. n.s. Finzi and Schlesinger
2003

Populus tremuloides 550 3 unfertilised n.s. ≠ n.s.3 n.s.3 Holmes et al. 2003
Betula papyrifera
Acer saccharum

Liquidambar 565 2 to 3 unfertilised n.s. n.s.3 n.s.3 Sinsabaugh et al. 2003

styraciflua
Crop Pr oduction
Oryza sativa A+200 1 to 2 8 to 9.4 g N m–2a–1 ≠ ↑ ↑ Inubushi et al. 2001

Note: Data was only included from field studies for which effects on microbial N were reported. Effects listed in parentheses are marginally significant.
1 Number of autotrophic ammonia oxidising bacteria.
2 Measured by pool dilution using 15N-labelled fertiliser; effect increases with time.
3 Gross rates measured by 15N pool dilution.
4 No effect on average across all years; there was a significant increase of C in 1 and a significant increase in N in 2 out of 6 years.
5 Nitrification potential measured in incubations.
6 Data in wetted soils as required for chloroform-fumigation-extraction method (Sparling et al. 1990).

21
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22 Agroecosystems in a Changing Climate

Soil P availability has been measured more directly in a number of studies,

mainly by bicarbonate extraction (Olsen-P). For example, Johnson et al. (2003)
reported reduced soil P availability in scrub oak forest exposed to elevated CO2 for
several years on some but not all sampling dates. However, no differences in P
collected on anion exchange resins were found in the same study.
The interpretation of soil phosphatase data is not straightforward for several
reasons. First, the production of extracellular phosphatases is supposedly induced
by a lack of available P. Increased phosphatase activities could increase mineralisa-
tion rates and ameliorate this condition; however, if the production of phosphatases
has been induced by a particularly low availability of P, mineralisation rates may
effectively still be reduced. Elevated phosphatase activities, therefore, can be inter-
preted either as signs of increased P mineralisation (because this is the purpose for
which they are produced) or of reduced supply to plants (because this is what induces
their production). Second, P can also be rendered plant-available by other avenues;
organic acids such as oxalate and citrate can effectively chelate phosphate and
improve its solubility, mobility, and plant availability. Third, measurement of soil P
pools by sequential extraction procedures yield indicators of plant-available forms
of P. However, low concentration can be a result of increased uptake by plants, or
the reason for low uptake, and active uptake mechanisms via organic acid exudation
and mycorrhizal networks are important and not accounted for by these extractions
for P (Rouhier and Read, 1998).
Enzyme activity and soil concentration measurements provide valuable infor-
mation, but in order to achieve a more conclusive understanding of CO2 effects on
P cycling, several methods would ideally be combined. These include a complete
assessment of ecosystem P pools (plants, soil microbial biomass, soil pool measure-
ments by sequential extraction procedures) and isotopic measurements of exchange
kinetics and microbial immobilisation (32P/33P).
Effects of elevated CO2 on S-cycling have not received much attention so far.
Mineralisation of S has been found to correlate with soil microbial activity in some
studies (e.g., Eriksen et al., 1995) and effects of elevated CO2 therefore appear
possible. Ebersberger et al. (2003) measured arylsulphatase activity in a calcareous
grassland and did not find any change under elevated CO2, but enzyme activities
may be poor indicators of S mineralisation rates. Supply of S has generally not been
a concern for agricultural production because atmospheric inputs from air pollution
are sufficient to cover plant demand, at least in industrialised areas. However, SO2
emission control has now led to a negative S balance in many agricultural soils (e.g.,
Knudsen and Pedersen, 1993). Plant S supply from mineralisation and potential
effects of elevated CO2 may become more important in the future.

2.2.3 PLANT TISSUE QUALITY

A significant part of plant biomass is exported as yield in many ecosystems (hay,
crop, animal biomass). The elemental composition of plant tissue is therefore a
critical determinant of the nutrient balance of the ecosystem. A shift in elemental
ratios under global change thus may shift the coupling of nutrient cycles and affect

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Climate Change Effects 23

nutrient limitations, also higher up the food chain. This may necessitate adaptations
in fertiliser use, and also alter the nutritional quality of plants.
Elevated CO2 affects plant tissue quality by several mechanisms:

1. Carbohydrate levels of green plant tissue increase, primarily in the form

of starch (Penuelas and Estiarte, 1998; Wong, 1990).
2. Leaf nitrogen concentrations decrease in many studies, mainly due to
increased carbohydrate contents and reduced amounts of photosynthetic
enzymes in C3 plants (e.g., Ainsworth et al., 2002; Rogers and Ellsworth,
2002; Sage et al., 1997; Sage, 2002; Seneweera et al., 2002; Stitt and
Krapp, 1999).
3. Leaf P decreases often less than leaf N, presumably because more P is
required in phosphorylated intermediates when photosynthetic capacity
increases under elevated CO2 (Ghannoum et al., Chapter 3, this volume;
and Gifford et al., 2000).
4. Allocation to secondary compounds may increase under elevated CO2 due
to reduced C and N limitation (Herms and Mattson, 1992; Penuelas and
Estiarte, 1998).

Not much is known about effects on elements other than N and P. What is the
experimental evidence that shifts in elemental composition occur under elevated
CO2, especially for elements other than N? In a search for patterns, I have compiled
data on pools and concentrations of nutrients in plants exposed to elevated CO2
(Table 2.2). I have mostly focused on multiyear field studies and excluded experi-
ments for which only N concentration was reported because the aim here is to assess
differential responses of mineral elements.
A remarkable finding is that foliar K concentrations decreased under elevated
CO2 in many studies conducted under relatively infertile conditions: Newbery et al.
(1995) reported decreases in shoot K concentration in Agrostis capillaris grown
under low K supply; specific root absorption, measured as 85Rb absorption of excised
roots, increased by over 100%, indicating increased demand for K. Decreased shoot
K concentrations were also reported by Schenk et al. (1997) for Lolium perenne/Tri-
folium repens swards. In trees, reductions in foliar K concentration were reported
for Quercus alba and Picea abies (Norby et al.,1986) and in mixed stands of Quercus
germinata/myrtifolia (Johnson et al., 2003). It is noteworthy that the decreases in K
were often larger than the decreases in N. These reductions in K translated into
reduced litter K concentrations where this was measured, and reductions in K again
were larger than reductions in N (mixed stands of Quercus germinata/myrtifolia:
Johnson et al., 2003; ombrotrophic Sphagnum bogs: Hoosbeek et al., 2002). This
decrease in foliar K occurs less frequently in well-fertilised systems, though there
are exceptions (e.g., cotton: Prior et al., 1998; Picea sitchensis: Murray et al., 1996).
It remains to be explored whether this decrease in K concentrations is ecologically
significant. Soils differ greatly with respect to K availability. Fine-textured soils have
generally larger exchange capacities, which prevent K leaching, and K is constantly
resupplied from mineral weathering. In these soils, K is generally not limiting.
However, sandy soils with low exchange capacity can result in significant K leaching,

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24
TABLE 2.2
Nutrient Pools and Concentrations in pLants Grown under Elevated CO2

CO2 Treatment
Type
of Conc. Nutrient
Ecosystem Study Soil (ppm) Duration Status Fraction Parameter N P K Mg Ca S Zn Mn Fe Cu B Si Reference

Grasslands
Calcareous field natural 600 6 yrs unfertilised shoot conc. decr.2 decr.2 – – – – – – – – – – Niklaus et al.
grassland soil N/P colimited 1998;
Niklaus &
Körner
2004

Tallgrass prairie field natural 2xA 3 yrs unfertilised shoots conc. decr.4 decr.4 – – – – – – – – – – Owensby et
soil al. 1993;
2 yrs litter conc. n.s. –47%* – – – – – – – – – – Kemp et al.
(Andropogon) 1994
2 yrs litter conc. n.s. n.s. – – – – – – – – – –
(Sorghastrum)
2 yrs litter (Poa) conc. –14%* n.s. – – – – – – – – – –

Agroecosystems in a Changing Climate

Lolium perenne/ pots natural 670 2 yrs full fertiliser at shoot conc. n.s. n.s. –18%** n.s. 10%** n.s. – – – – – – Schenk et al.
Trifolium soil 200 kgN /ha shoot pool 22%** 20%* 3% 22%** 37%** 20%** – – – – – – 1997
repens swards

Agrostis capillaris pots 680 43 d fertilised shoot pool +122%* +193%* +135%* +82%* +175%* – – – – – – – Baxter et al.
Poa alpina in 105 d pool +13% +23%* +15% +25%* –48%* – – – – – – – 1994
Festuca vivipara OTC 189 d pool –73%* –48%* –38%* –37%* –45%* – – – – – – –
Agrostis capillaris 43 d conc. n.s. n.s. n.s. n.s. n.s. – – – – – – –
Poa alpina 105 d conc. n.s. n.s. n.s. n.s. n.s. – – – – – – –
Festuca vivipara 189 d conc. n.s. n.s. n.s. n.s. n.s. – – – – – – –

Agrostis pots sand A+250 23 wks Modified shoot conc. –9% –13% –38%** – – – – – – – – – Newbery et
capillaris hoadland pool +8% –3% –23% – – – – – – – – – al. 1995
solution with roots (excised) absorption n.s. n.s.
+100% – – – – – – – – –
variable con- ***
centrations
of NPK
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Climate Change Effects

Agricultural Crops
Oryza sativa field rice paddy A+200 123 d NPK–fertiliser grain conc. –10%* –4% –9%** –6% –1% – – – – – – +4% Yamakawa et
al. 2003

Triticum pots sieved 700 116d NPKCa– whole plant5 conc. –4%** 3% –11%*** – – – – – – – – – van Vuuren et
aestivum arable fertiliser pool 5% +12%* –3%# – – – – – – – – – al. 1997

Gossypium field arable soil 550 150 d full fertiliser leaves conc. –7%# +2% +4%* +5%# –3% – +15%# +3% +12%# +10%# – – Prior et al.
hirsutum whole plant incl. conc. –11%* –2% –7%# –6%# –14%* – –3%# –9%# –8%# –9%# – – 1989
root
leaves pool +7%# +17%# +19%# +22% +12% – +33%# +20% +33% +28%# – –
whole plant incl. pool +21% +33%* +26%# +27%# +17% – +31%* +24% +26% +24%# – –
root

Trees
Quercus rubra OTC natural 700 2 yrs natural leaves conc. –4%* +8% –15%* +13% –3%* +6% – –25%* – – – – Le Thiec et
(3 yr old) soil al. 1995
Picea abies OTC natural natural needles conc. –4%* +21% –6%* –36%* –28%* +8% – –30%* – – – –
(5 yr old) soil

Picea sitchensis pots in artificial A+350 3 yrs full fertiliser current year conc. –15%* n.s. n.s. – – – – – – – – – Murray 1996
branch needles
OTC mixture previous year conc. –7% n.s. n.s. – – – – – – – – –
branch needles

Citrus aurantium OTC natural A+300 4 yrs full fertiliser leaves conc. decr. n.s. decr. n.s. decr. n.s. n.s. decr. n.s. n.s. n.s. – Gries et al.
soil 1993
Quercus field natural A+350 5 yrs unfertilised shoot (Q. pool 13% 18% 3% 44% 33% 10% 53% 58% 18% 60% 15% – Johnson et al.
germinata/ soil myrtifolia + 2003
Quercus Q. germinata)
myrtifolia shoot (Q. conc. –17%** –14%** –23%** +0% –4% –23% ** +6% +13% –11% +14% –18% –
stands myrtifolia +
Q. germinata)
litterfall conc. +1% +11% –9%** +8% +8% –20%** –3% +53% –26% –63% –22% –
*** ***
standing litter conc. +4% –20% –10%** +0% +10% –16% ** +30%* +37% +8% +67% –18%* –
**

25
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26
TABLE 2.2 (CONTINUED)
Nutrient Pools and Concentrations in pLants Grown under Elevated CO2

CO2 Treatment
Type
of Conc. Nutrient
Ecosystem Study Soil (ppm) Duration Status Fraction Parameter N P K Mg Ca S Zn Mn Fe Cu B Si Reference

Fagus sylvatica/ OTC acidic loam shoots (Fagus + pool –2% –1% +4% –3% +5% +2% +8% +6% +12% – – – Hagedorn et
Picea abies soil Picea) al. 2002 .
Fagus sylvatica/ calcareous shoots (Fagus + pool +13% +63% +38% +30% +29% +17% +56% +32% +60% – – –
Picea abies sand Picea) ** *** *** *** *** *** *** ** **
Fagus sylvatica acidic loam foliage conc. –7% –6% 3% –9% 4% –10% 0% 15% 4% – – – Hagedorn,
soil pers. comm
calcareous foliage conc. –10% 18% 1% –14% 1% –7% 29% –24% –11% – – –
sand
Picea abies acidic loam foliage conc. –19% –18% –17% –12% 15% –13% 15% 6% –10% – – –
soil
calcareous foliage conc. –12% 5% –6% –6% 9% –7% 28% –3% 1% – – –
sand

Pinus ponderosa OTC natural 700 2–3 yrs6 unfertiliser, needles conc. –8%# –4%# –12% –17% –1% –9%# –13% 5% +22%# –7% –11% – Walker et al.
soil low and high 2000

Agroecosystems in a Changing Climate

(NH4)2SO4
application

Other Ecosystems
Ombrotrophic field natural 560 3 yrs unfertilised Sphagnum litter conc. –4% –10% –14%* – – – – – – – – – Hoosbeek et
bogs (4 sites soil Eriophorum leaf conc. –13%* –3% –8%** – – – – – – – – – al. 2003
across Europe) litter
Eriophorum conc. –9% +3% +23% – – – – – – – – –
root litter

Note: Only studies also reporting nutrients other than N were included, and preference was given to multiyear field experiments. When studies contained factorial treatments, responses were calculated for each treatment combination and average responses
are listed; the same was done for multiyear data sets. Significant effects are indicated by * P < 0.05, ** P < 0.01 and, *** P < 0.001, is used to indicate P < 0.1 and is also used when significances are not obvious, for example, for the average response
of multiyear data sets where significant effects were found in some but not all years. Refer to the original publications for details.
1 There were transient responses that vanished at end of experiment.
2 Decrease in P was smaller than decrease in N.
3 Measured with 15N, 32P, and 85Rb.
4 Effect on [N] not present in all species at all dates but larger than on [P], which decreased in 1 species and 1 out of 3 years.
5 Average of wet and dry treatment shown.
6 Total study duration was 5 years; average effects across 2nd and 3rd year of treatment and all CO2 and N fertiliser levels given here; # indicates significant effect in at least one year.
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Climate Change Effects 27

especially in high rainfall areas. For example, rangelands of western Australia often
exhibit severe K limitation (Bolland et al., 2002; Cox, 1973) and effects of elevated
CO2 on K contents of grass, therefore, may be important. Moreover, elevated CO2
may affect leaching rates and associated K losses in these ecosystems.
Data for all other nutrients is far too limited to attempt any generalisations;
remarkable effects are the decrease in leaf S and B concentrations in the mixed
Quercus stands investigated by Johnson et al. (2003) and decreased Mn concentra-
tions in Quercus alba and Picea abies (Norby et al., 1986).

2.2.4 LEACHING AND VOLATILISATION

Higher soil moisture has been reported in many ecosystems exposed to elevated CO2
(Morgan et al., 2004b) and this will favour the leaching of nutrients because more
water will drain when saturation is exceeded. However, other factors that may affect
leaching have also been found to change under elevated CO2: Soil aggregation was
found to increase (Rillig et al., 1999) or decrease (Niklaus et al., 2003, 2001a) under
elevated CO2, and Newton et al. (2004) reported reductions in soil hydrophobicity.
Dry soils also tend to shrink and form cracks, a phenomenon that may be less
pronounced under elevated CO2 because soils might be moister. Increased root
production under elevated CO2 might be another factor reducing crack formation.
Finally, detritus is an important determinant of soil hydrophobic properties. What
the combined effect of all these changes on preferential flowpaths and water retention
remains elusive.
Direct evidence for nutrient leaching in elevated CO2 studies is both scarce and
controversial. Körner and Arnone (1992), for example, have reported increased NO 3−
leaching from tropical communities, while Torbert et al. (1996) found decreased
NO 3− concentrations below the rooting zone. Niklaus et al. (2001b) observed strong
and persistent reductions in soil NO 3− concentrations in calcareous grassland com-
munities exposed to elevated CO2 for several years, but it is not clear how much
NO 3− was leached from this system. Soil NO 3− concentrations are regulated by many
interacting processes, including nitrification and denitrification, immobilisation of
NH +4 and NO 3− by soil microbes, and rooting patterns and root uptake kinetics
for NH +4 and NO 3− . There is also evidence that plant secondary compounds (terpenes
and tannins) can inhibit nitrification (Northup et al., 1998, 1995; Olson and Reiners,
1983; Reiners, 1981; Rice and Pancholy, 1973, 1974; White, 1986, 1988; White and
Gosz, 1987) and this process may be more important at elevated CO2. Therefore,
predictions are difficult, but a principal control is certainly plant uptake of mineral
N, which will reduce NH +4 available for nitrification or remove the NO 3− produced.
Andrews and Schlesinger (2001) studied a forest site exposed to elevated CO2
and reported increased soil CO2 concentrations, which were caused by higher plant
and microbial respiration. These increased soil CO2 concentrations accelerated soil
acidification and mineral weathering, and there was evidence of increased leaching
of bicarbonate base cations. This is another mechanism by which extra nutrient
leaching losses may occur under elevated CO2.

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28 Agroecosystems in a Changing Climate

2.3 EFFECTS OF ELEVATED TEMPERATURE AND

PRECIPITATION ON NUTRIENT CYCLING
Elevated CO2, together with other radiatively active atmospheric gases, will lead to
higher global mean temperatures and altered precipitation patterns (IPCC, 2001).
These climatic drivers differ in an important aspect from the increase in atmospheric
CO2. For any practical purpose considered here, atmospheric CO2 concentrations
are globally and temporally uniform. Therefore, effects found in different studies
are relatively easy to compare. In contrast, even in the absence of global change,
temperature and humidity change constantly in any ecosystem, even at the time scale
of hours, and the ecosystem is constantly responding to these alterations. In the long
term, temperature and precipitation define large-scale vegetation distribution and
biomes; in the short term, however, ecosystems are very persistent to even extreme
episodes such as droughts or exceptionally cold spells.
Many ecosystem processes exhibit a pronounced temperature optimum, and
ecosystems often operate above or below this optimum — this may even change in
the course of the day or season. Consequently, a superimposed experimental or
“natural” future warming may have positive or negative effects, depending on the
ecosystem studied and the actual climatic conditions prevailing. This greatly exac-
erbates difficulties in the attempt to identify common response patterns across
studies.
Several avenues have been pursued to study warming effects on ecosystems. A
number of studies have capitalised on existing geographical gradients in temperature
(e.g., Jenny, 1980; Raich and Schlesinger, 1992; Saleska et al., 1999; Trumbore et
al., 1996) and on natural inter-annual variability (e.g., Chapin et al., 1995; Oechel
et al., 2000). While this approach is largely correlative, it has the advantage that it
addresses changes over longer time scales and largely avoids experimental artifacts.
Alternatively, ecosystem temperature has been manipulated by heating soils with
buried electric cables (e.g., McHale et al., 1998; Peterjohn et al., 1993; Rustad and
Fernandez, 1998), with infrared lamps suspended over vegetation (e.g., Saleska et
al., 1999), and by using passive open-top-like structures that increase temperatures
by acting as a small greenhouse (e.g., Arft et al., 1999; Robinson et al., 1995). While
these manipulative studies allow us to causally attribute detected effects to the
warming treatment, they are often associated with experimental problems. For exam-
ple, infrared heating, as used in many studies, is characterised by largely lacking a
convective heating component, and buried cables may lead to heterogenous temper-
ature distribution, especially in dry soils.
Effects of soil moisture and temperature on microbial activities often are inter-
active because moisture optima become more narrow at elevated temperatures (Fig-
ure 2.3). A good example is the data by Goncalves and Carlyle (1994) who studied
N mineralisation in vitro at different temperatures and soil moisture contents. The
reduction in mineralisation rate when reducing soil moisture from 60% to 10% of
field capacity was much larger relatively at 25ºC (≈70%) than at 5ºC (≈50%).
Warming, both experimentally and naturally, is often accompanied by increased
evapotranspiration and, therefore, decreased soil moisture and plant water availabil-
ity (e.g., Luo et al., 2001; Saleska et al., 1999), which results in a further reduction

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Climate Change Effects 29

FIGURE 2.3 Schematic representation of interactive effects of temperature and moisture on

microbial activity. Arrows show hypothetical increases in microbial activity as caused by a
temperature rise. Three different scenarios, a through c, with respect to soil moisture are
shown. At the low temperature, soil moisture is optimal in all three scenarios and microbial
activity equal. Arrows a and b show effects of temperature increases without concomitant
change in soil moisture; note that the effect size in scenario b is smaller than in scenario a
because the optimal soil moisture range becomes narrower at elevated temperature. Scenario
c shows the situation that is typical of many manipulative warming studies in which temper-
ature increases are accompanied by a concomitant drying of soils or litter layer. In this case,
activities are further reduced by the actual drying of soils. (Drawing based on data by Bunnell
and Tate [1974] as shown in Paul, E.A. and Clark, F.E. [1996]; scenarios added by author.)

of microbial activity (arrow c in Figure 2.3). As discussed, elevated CO2 also affects
water relations (though in the opposite direction) and this is clearly a very important
mechanism of responses to elevated CO2. In warming studies, however, this indirect
effect via alterations of water relations is even more important. It is often very
difficult to separate these two factors, and confounding interpretations may be the
result.
A further point to consider is that soil temperatures are determined by a number
of processes, including radiative heating, convective energy exchange with air layers
above the soil, and by heat conductance within soils (Figure 2.4). As a consequence,
soil temperatures do not necessarily track ambient air temperatures. On the contrary,
several whole-ecosystem studies have shown that increased ambient air temperatures
can actually result in decreased soil temperatures (Coulson et al., 1993; Robinson
et al., 1995; Wookey et al., 1993). While this appears to be counterintuitive at first
glance, the phenomenon becomes understandable when considering that increased
biomass of plants at elevated ambient temperatures can effectively insulate soils
from solar radiation; furthermore, taller-stature species will absorb solar radiation
farther off the ground and convective heating of soils will effectively be reduced (T1
and T2 in Figure 2.4). During the cold season, a related temperature reversal can
happen as a consequence of snowpack (Groffman et al., 2001a). A warmer climate
is likely to be associated with less frequent and shorter snow cover. Because snow
very effectively insulates soil from convective and radiative heat loss, more frequent

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30 Agroecosystems in a Changing Climate

FIGURE 2.4 Schematic representation of vegetation effects on soil temperature. Incident

radiation is converted to heat where it intercepts plant canopies. In taller plant canopies, this
will happen farther off the ground and convective heating of soils will be smaller than in low-
stature canopies. Soils under tall canopies, therefore, will be cooler (T1) than under small
canopies (T2), which will be cooler than bare ground (T3). Conductive cooling of top soils
by cooler subsurface soils will depend on heat conductivity and heat capacity of soils, which
are strongly influenced by moisture contents. (Latent heat fluxes due to evapotranspiration
are not shown.)

soil freezing will occur. This is all the more important because root and microbial
processes are still active at temperatures near or even below 0°C, and because
freeze–thawing cycles can dramatically affect these processes (Groffman et al.,
2001b).

2.3.1 NET PRIMARY PRODUCTION, DECOMPOSITION, AND THE

CARBON AND NITROGEN BALANCE OF ECOSYSTEMS
Soil organic matter levels are determined by the balance between primary production
and decomposition. An important question is whether productivity and decomposi-
tion respond similarly to climate change; if they do not, soil organic matter pools
will change, resulting in concomitant long-term release or immobilisation of nutri-
ents. It has been hypothesised that soil organic matter decomposition will increase
more strongly at elevated temperatures than net primary productivity will, resulting
in an imbalance between inputs and outputs to the ecosystem and, therefore, in soil
organic matter losses. But what is the available evidence to support this claim?
Plant growth is limited by temperature in many ecosystems (Churkina and
Running, 1998). Several growth-determining processes (e.g., leaf expansion rates or
photosynthesis) are directly dependent on temperature, and the length of the growing
season may also increase in a warmer climate. Soil microbial respiration, at least
when measured on isolated samples and, in the short term, generally increases with
temperature, often exponentially, at least when soil moisture is optimal and temper-
atures are not too high (cf. Figure 2.3). When both production and decomposition
are considered separately, one might conclude that the temperature dependence of
respiration is stronger (i.e., exponential), and that a net decomposition of new organic
matter will happen at elevated temperatures.

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Climate Change Effects 31

However, under steady-state conditions, long-term plant productivity and decom-

position are approximately balanced. If there is a disequilibrium, substrate available
for decomposition will increase or decrease until equilibrium is reached again
(decomposition generally follows first-order kinetics). From that, one might conclude
that respiration is determined by net primary productivity and this notion is often
found in the literature (e.g., Kirschbaum, 1995). However, it ignores feedback
mechanisms of decomposition on primary productivity, for example via the nitrogen
cycle. If primary production is N-limited and N availability controlled by decom-
position, one might as well contend that decomposition controls primary productiv-
ity; this shows that the involved feedback mechanisms clearly are more complex
and multidirectional.
Natural temperature gradients offer an opportunity to avoid circularity and to
study production and decomposition in an integrated context. Jenny (1980), in his
now classical work, compared tall grass prairie hay yields across a north–south
transect through the midwestern United States. Yields remained very constant,
although the gradient covered a mean annual temperature range from below 5ºC
to 20ºC. Kirschbaum (2000) argued that there probably was increased moisture
limitation at the warmer sites, and that Jenny might have found a positive effect
of warming on productivity if water had not been limiting. Jenny also investigated
a similar gradient (5 to 14ºC; Canada to Texas) in the more western short grass
belt, but again no temperature dependency of productivity was found. Post et al.
(1982) analysed the dependence of soil carbon content on climate. Climate and
corresponding life zones were classified according to temperature, precipitation
and potential evapotranspiration (PET) (Holdridge classification; Holdridge,
1947). Soil carbon stores were negatively correlated with temperature, but posi-
tively correlated with precipitation and precipitation:PET (which is an index of
water availability). However, Post et al. (1982) found a large variation of soil C
within individual life zones, indicating that local conditions and other factors were
also very important determinants. Townsend et al. (1995) studied soil organic
matter dynamics and respiration rates along an elevation gradient covering ≈10ºC
on the island of Hawaii; soil organic matter contents increased with altitude (i.e.,
decreased with temperature), and mean annual respiration rates showed the oppo-
site effect. Other gradient studies with similar results exist, and generally suggest
that soil organic matter might decrease in a warmer climate. However, there is
also evidence for the opposite. For example, Liski and Westman (1997) studied
soil carbon contents along a climosequence across Finland spanning 800 to
1300ºC·days*; topography, parent material, and soil age were standardised care-
fully. Two site types differing in the type of coniferous forest were sampled: Pinus
sylvestris/Calluna sites had low productivity, whereas Picea abies/Myrtillus sites
had higher productivity. Soils were sampled to 1 meter depth and organic and
mineral soil C analysed separately; at the warmer end of the sequence, mineral
soil C was 57% (low productivity sites) and 28% (high productivity sites) higher
than at the cool end, and temperature accounted for 33 to 41% of the observed
variation in soil C. Organic soil C pools did not vary with temperature.

* A threshold of +5ºC was used.

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32 Agroecosystems in a Changing Climate

Matters are more complicated with respect to soil nitrogen because soil N does
not change in concert with soil organic C. Post et al. (1985), in their global survey
of soil profiles, found that soil nitrogen pools followed similar trends as they had
previously found for C (i.e., N was negatively correlated with temperature and
positively correlated with precipitation and precipitation:PET). However, soil C:N
ranged from <10 in tropical deserts, over intermediate values in the temperate zone
(10 to 20), to >20 in wet forests. Interestingly, the reason why wet forests had high
C:N depended on temperature was that in the wet tropics, C:N was high because
SOM (soil organic matter) was stored as relatively recalcitrant humic substances in
advanced stages of decay; in wet tundra C:N was high because decomposition was
slow. Jenny (1980) analysed N pools in the top 20 centimetres of soils collected in
the North American Great Plains; N pools decreased with temperature. However,
Jenny found a strong dependence on soil moisture, with decreasing nitrogen contents
at elevated mean annual NSQ.* Jenny also reported decreasing mineral soil C:N
along an altitude transect at Kaiser Pass (near Fresno, CA) along which temperature
decreased from 16 to 6°C and precipitation doubled from 450 to 900 mm a–1.
Manipulative soil warming experiments have shown increased soil respiration at
elevated temperature (e.g., Peterjohn et al., 1994); an important insight, however, was
that respiration often acclimates relatively quickly to warmer conditions, that is, res-
piration responses decrease with time (Luo et al., 2001; McHale et al., 1998; Melillo
et al., 2002). Soil respiration–temperature dependencies as found in short-term warm-
ing studies, therefore, cannot be extrapolated to longer time frames, or an overestima-
tion of soil organic matter losses will result (Eliasson et al., 2005). Simulation studies
also indicate that what is experimentally observed as acclimation may, in fact, not be
an acclimation of microbial communities but a depletion in young organic matter
fractions (Eliasson et al., 2005). New inputs of plant organic matter decompose quicker
than old soil organic fractions; this is evidenced in radiocarbon age of soil respiration,
which is much smaller than the age of bulk soil C (Trumbore, 2000). At low temper-
atures, fresh (active) soil organic matter turns over more slowly than under warmer
conditions (Trumbore et al., 1996); as a consequence, acclimation responses to warm-
ing should be smaller and slower in cold than in warmer climates, and this indeed was
found in several studies (e.g., Luo et al., 2001; Oechel et al., 2000).
At the ecosystem level, a number of important feedback mechanisms can alter soil
organic matter decomposition, sometimes in rather unexpected ways. As already noted,
several studies have shown that increased ambient air temperatures can result in
decreased soil temperatures (Figure 2.4 and Coulson et al., 1993; Robinson et al.,
1995; Wookey et al., 1993). Another important ecosystem-level feedback mechanism
is that plant community composition may change in response to the warming treatment;
a good example is the study by Saleska et al., (1999), which showed that warming of
a montane meadow induced soil drying, which in turn led to a shift from productive
to less productive but more drought-tolerant species. The warmed plots had lower soil
respiration rates than control plots because soils were drier, but also because the input
of decomposable material to soils was lowered due to the reduced productivity of the

* NSQ equals precipitation divided by the absolute H2O saturation deficit of air, and correlates with
precipitation:potential evaporation.

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Climate Change Effects 33

drought-tolerant species. Such species responses can be slow: Chapin et al. (1995)
studied tussock tundra under increased temperatures and found that responses after 3
years were poor predictors of longer-term (9 years) changes in community composi-
tion. Responses after nine years showed closer resemblance to patterns of vegetation
distribution along natural environmental gradients.
Finally, increased temperatures may lead to increased decomposition rates and
thus increased nutrient availability; plant productivity, therefore, may increase if
nutrients are limiting. Because C:N of plant material is, on average, larger than C:N
of soils, soil organic matter decomposition may effectively lead to ecosystem-level
C sequestration. However, if a substantial fraction of the nutrients mineralised is
lost, for example, by leaching, the ecosystem will become a net source of CO2 (e.g.,
Melillo et al., 2002; Shaver et al., 1992).
To conclude, predicting effects of warming on soil organic matter levels is very
difficult. Correlative studies have the advantage that they assess responses near
steady state, and thus also account for slow soil and vegetation adjustments; they
may, however, miss transient effects that are real, that is, those that occur over
decades to a century, which is the time frame over which temperature increases in
the range of several degrees are expected. The other extreme involves warming
studies conducted with isolated soil samples; it appears very dangerous to extrapolate
from such data because plant production and microbial decomposition are intimately
intertwined at many levels (cf. previous discussion of ecosystem-level feedback),
and these ecological mechanisms are effectively eliminated by separating soils from
plants and natural climate variability. Laboratory incubations, however, are very
helpful when combined with ecosystem-level field studies because they allow for a
better process-level understanding of effects observed in the field. Long-term whole-
ecosystem warming experiments (Chapin et al., 1995, e.g.,) or reciprocal transplant
studies along natural environmental gradients (e.g., Ineson et al., 1998) are probably
a good middle ground in that they allow for important ecological feedback mecha-
nisms (that may currently be unknown), can be combined with additional treatments
(e.g., manipulations of soil moisture), and allow us to follow the trajectory along
which the ecosystem adjusts to the new climate.

2.3.2 NUTRIENT IMMOBILISATION AND MINERALISATION

As discussed, increased temperatures might change organic matter contents and
associated nutrient pools in the long term; but what is the available direct evidence
that plant-available nutrients (resulting from net mineralisation) change in the short
term, and what is known about the underlying processes?
The strong temperature dependency of organic matter decomposition may accel-
erate N mineralisation in a warmer climate. At the same time, a number of processes
may affect mineralisation in the opposite direction:

1. Warming often results in decreased soil moisture, at least in manipulative

experiments, which could decrease mineralisation rates (e.g., Robinson
et al., 1995).

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34 Agroecosystems in a Changing Climate

2. Some plant species respond to increased temperature with increased lig-

nification, which could reduce decomposability of litter and nutrient
release rates.
3. The composition of plant communities may shift in response to warming,
with more lignified species prevailing at elevated temperatures.
4. Increased biomass production may cool the litter layer and top soil (Figure
2.4), which may reduce decomposition rates.

Net mineralisation is the outcome of gross mineralisation and immobilisation,

each of which is controlled by a complex of factors. A meta-analysis by Rustad et
al., (2001) revealed, on average, an increase in N mineralisation rates in response
to elevated temperatures, although there were exceptions. Nadelhoffer et al. (1991)
incubated different soils from Alaskan tundra at 3, 9, and 15ºC, and periodically
leached soils to measure NH +4 , NO 3− , and PO 3− 4 in the leachates collected. An
important finding was that the increase in N mineralisation was much greater from
9 to 15ºC than from 3 to 9ºC, despite the marked increase in temperature dependency,
which is often found at low temperatures (Kirschbaum, 1995). Soil respiration
behaved similarly, and the authors attributed this unexpected temperature depen-
dence to the different temperature sensitivities of the enzymes involved in cellulose
degradation. Interestingly, the release of PO 3− 4 was often reduced at elevated tem-
peratures. In some soils, PO 3−
4 release was 5 to 10 times greater at 3ºC than at 9ºC
and 15ºC, and Nadelhoffer et al. (1991) concluded that P immobilisation increases
more rapidly than gross P mineralisation, and that this might result in increased P
limitation under warmer conditions.
Tscherko et al. (2001) measured soil enzyme activities in microcosms subjected
to experimental warming. Microbial biomass N did not change in this study, and
the activities of soil enzymes related to N mineralisation remained mostly unaltered
(with the exception of arginine deaminase, which was increased).
Rustad and Fernandez (1998) studied the decomposition of red spruce and red
maple litter in temperate forest floor electrically warmed by 5ºC. All litter had grown
under ambient conditions, so that the initial quality did not differ between treatments.
Interestingly, both litter types decomposed quicker in the heated treatments, but
decomposition (mass loss) was accelerated during the first 6 months in maple, and
during the 18- to 30-month period in spruce. K and B were released quicker in
maple, while Ca, Mg, K, and Zn were released more quickly in spruce. No differ-
ences were found for the other elements (Cu, Mn, N, Al, Fe). This study nicely
demonstrates that warming effects on decomposition are time and element specific.
Chapin et al. (1995) increased ambient temperatures by 3.5ºC over tundra veg-
etation plots for 9 years with the help of plastic tents. Soil NH +4 and PO 3−4 concen-
trations were increased in the heated treatment and increased amounts of NO 3− were
collected on soil-buried ion exchange resins. N and P availability actually were
increased during winter after the greenhouses had been removed from heated plots,
and Chapin et al. therefore concluded that changes in mineralisation were not just
due to direct warming effects on soil processes, but rather a consequence of changes
in species composition and litter quality. Total N and P pools in vegetation did not
change at elevated temperature when plots were not fertilised. However, mosses and

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*** START OF THE PROJECT GUTENBERG EBOOK THE HILL OF

ADVENTURE ***
“Things have broken loose quicker than we
thought”
 THE HILL OF ADVENTURE
BY ADAIR ALDON
Author of “At The Sign of the Two Heroes,” etc.

ILLUSTRATED BY
J. CLINTON SHEPHERD

NEW YORK
THE CENTURY CO.
1922
Copyright, 1922, by
The Century Co.

PRINTED IN U. S. A.
 LIST OF ILLUSTRATIONS
“Things have broken loose quicker than we thought”
“It is you who do not understand,” he returned gravely
He had only to lift his voice, and the long spell would be broken
Beatrice found herself telling what had happened
THE HILL OF ADVENTURE
 CHAPTER I
GRAY CLOUD MOUNTAIN

It was with feelings of doubt that were not very far from dismay that
Beatrice Deems watched her new acquaintance, Dan O’Leary, saddle
her recently acquired horse. She had ridden before, of course, in the
tan-bark ring of the riding-school or on shady bridle-paths in the
park, always on well-broken steeds whose beauty and grooming
were equaled only by their good manners. But now, as she stood in
her short khaki riding-skirt and her high boots, waiting outside the
great dilapidated shed that, in this little Montana town, did duty as a
livery-stable, she was beginning to wonder whether she really knew
anything about horses at all. Certainly she had never thought of
riding anything like this plunging creature who stood straight up on
his hind legs one moment, then dropped to his forefeet and stood on
them in turn, with the ease of a circus performer.
She had spent only two days in Ely, the little town planted beside
Broken Bow Creek, in the foot-hills of the Rocky Mountains. At first
she had thought that the village, with its scattered box-like houses
and dusty, shadeless street, was disappointingly unlike the West of
the pictures-books and the movies. The antics of her new horse,
however, were disturbingly like what she had witnessed in Wild West
shows.
“’Name’s Buck,” volunteered the man who was struggling with the
saddle, and added, though in a tone that seemed to indicate the
explanation as quite unnecessary, “It’s on account of his color, you
know.”
“Oh!” returned Beatrice, a little blankly. For the life of her, she
could think of nothing else to say. She had yet to learn that all
Western ponies of that golden buckskin shade of coat bear the same
name. At the moment she was tempted to believe that the title had
something to do with the way in which the horse was humping his
back like a gigantic cat and jumping up and down on his nimble
white forefeet.
“Your father went out on the range and chose the horse himself
when he was out here getting your house ready,” Dan went on. “He
couldn’t have found another pony in the valley that could go like this
one.”
“Did he—did he try him?” Beatrice wished to know.
Her feelings in the matter were oddly mixed, for she dreaded the
moment when she must actually mount to the big, unfamiliar saddle,
and yet she was all on fire to try the horse’s speed.
“No, he didn’t try him,” was the answer. “He just said he wanted a
safe horse for his daughter, liked the looks of this one—and well he
might,—and took my word for it that the horse would suit and would
go like greased lightning, besides. There, now, the saddle’s firm. You
mustn’t think anything of the way he acts when you pull up the
cinch; they all do that!”
For all her misgivings, Beatrice was no coward. She stepped
forward, discovered in one violent second that a Western pony sets
off the moment he feels the rider’s weight on the stirrup, then flung
herself, somehow, into the saddle and was away.
“I did not do that very well,” she was thinking. “Another time—oh,
oh!”
For her very thought was interrupted by the sudden rush of
wordless delight as the horse beneath her stretched himself to that
long easy lope that is like nothing else in the world. The fresh
mountain wind, sweeping down from the clean, high peaks above,
sang in her ears; the stony road swung past below; the motion was
as easy as a rocking-chair but seemed as swift as thought itself.
Motoring she had always loved, but she confessed with sudden
disloyalty that it was bumpy business compared to the measured
swaying of this living creature between her knees. Buck’s personal
prejudices seemed, indeed, to be directed solely against the cinching
of the saddle. That process once over he was as eager and happy as
she to clatter across the bridge, pass the last of the ugly little houses
and the high-fronted store buildings, and turn his white-blazed face
toward the mounting trail that led out of the valley.
Beatrice drew rein when they had breasted the first rise, and
paused a moment to look back. The houses strewn haphazard
across the slope below her made more of a town than she had
thought. There was the packing-box railroad station where she and
her sister, Nancy, and their Aunt Anna had arrived so recently; there
was the house where they were living, a little larger than the others,
but square, hideous, and unshaded like the rest.
“We mustn’t care for architecture,” Nancy had said when they first
surveyed their dwelling rather ruefully, “when the Rocky Mountains
begin in our back yard.”
There was also the winding stream with its abrupt bend that
warranted the title of Broken Bow Creek, a mere trickle of water just
now, in that wide, dry valley down which the thin line of the railroad
stretched away, with the straight parallel of the rails seeming to
bend and quiver in the hot clearness of the sunshine. South of the
town was a portion of Ely that she had not seen before, a group of
warehouses, some office buildings, and a huddle of workmen’s bunk-
houses. She could see the cobweb lines of temporary railroad, a
steam-shovel moving on a flat-car, and innumerable men toiling like
black ants along the sides of the raw cut that had been made in the
red soil of the valley.
“That must be the irrigation ditch that Dan O’Leary was telling us
about,” she reflected. “How hot it looks down there! I did not dream
they had so many men. And how clear the air is! Oh, surely, surely,
Aunt Anna will get well here as fast as we hope!”
The wind lifted Buck’s yellow mane and her own brown hair, while
the horse pawed the stony ground impatiently. She let him go on,
for she was in truth as eager as he. This was the first day that she
had found time to go far from their own house, and she had now a
most fascinating goal before her. What girl of sixteen would not feel
excited over the prospect of exploring a tract of mountainside woods
of which she was sole owner?
 Beatrice had never quite understood how her father had come to
purchase that stretch of land above Ely; she had not, indeed,
thought to ask. She had come into his study one Sunday morning
when he was going over his papers and had surprised him with the
announcement that she was sixteen that day. Having no other
present ready, he had brought out some dusty title-deeds and had
made them over to her.
“It will never be of the least use to you, my dear,” he said, “so do
not consider it much of a present. Twenty-three acres with timber,
cabin, and a waterfall, so the description reads, but you must not
think they are worth anything. I have never seen the place myself.”
She had believed that it was on account of this talk about Ely that
they thought of the town again when the doctors had prescribed for
Aunt Anna “a change of climate—some dry, bracing place in the
West.” She was their only aunt, Mr. Deems’s younger sister, and she
had cared for his household ever since the death of the two girls’
mother years ago. She was a slim, frail person of indomitable spirit,
and had begun to look as though she were far more spirit than body
ever since the influenza epidemic had swept through the family.
Beatrice had always thought that going to Ely was her own
suggestion, though she could not deny that it was Aunt Anna who
had carried the plan through in the face of some rather
unaccountable opposition from her father. Mr. Deems had finally
given in, and had made a flying trip to Ely to be sure that the air and
climate were what they wanted, to choose a house, engage a
Chinese cook, and make all preparations for a summer’s stay for his
sister and the two girls.
“I did not have time to visit your estate on the hill, Beatrice,” he
said on his return. “You will have to explore it yourself. Dan O’Leary
has charge of it and said he rented it to some engineers who were
surveying the mountain, but it is unoccupied now. The place may
prove to be a good picnic ground but I fear it has no other
possibilities.”
He might say what he chose, Beatrice was thinking, but he could
not destroy her eagerness to see the place. The trail ran crookedly
upward before her, disappeared in some dense pine woods, then
slanted across the spur of the mountain and vanished again. Higher
above rose the bare, rocky slopes of the tall peak that dominated
the whole valley, Gray Cloud Mountain, on one of whose lower,
rugged shoulders lay her land and her cabin. After climbing for a
quarter of a mile, she was obliged to hesitate at a fork in the way,
uncertain which of the steep paths she was to take.
A little cottage clung to the bare hillside by the road—a shabby
place with no paint and a patched roof. The door was swinging open
as she passed and a man just going in, a short-set, foreign-looking
person, who scowled at her over his shoulder when she asked the
way.
“That one,” he said briefly, pointing to the right-hand fork and
speaking with a heavy foreign accent. “Up toward John Herrick’s
house, only not so far.”
He went in and shut the door abruptly. Beatrice could hear his
voice inside calling roughly, “Christina, Christina!”
He had a roll of large papers in his hand, posters that he had
evidently been putting up along the way, for she had observed them
on trees and fence-posts nearer town. They seemed to announce a
meeting of some sort, with English words at the top and odd foreign
printing at the bottom in more than one language. She had felt a hot
flash of indignant anger at the man’s surly tone, but in a moment
she had forgotten him completely, as she and Buck went scrambling
up the steep and difficult road.
She came at last to a tiny bridge. Broken Bow Creek, which was
little more than a series of pools in the parched stream-bed in the
valley, was here a singing rivulet, flowing below the rude crossing
amid a group of silvery aspen-trees. At the left of the trail she could
see a gate, a set of bars hung between two rough posts. It was with
a beating heart that she dismounted to take them down for Buck to
pass. Once inside she would be on her own ground.
The agility of a mountain-bred pony was so new to her that she
was much astonished, after she had removed two of the bars, to
have Buck step over the remaining three as neatly as a dog would
have done. She slipped into the saddle again, making a greater
success than at the first attempt, and followed the nearly invisible
path. The huge straight pine-trees stood in uneven ranks all about
her, their branches interweaving overhead, the ground covered with
their red-brown needles that muffled the sound of the horse’s hoofs.
Up they went, with the splash of falling water sounding louder and
ever louder. Here at last was the place she sought, a square, sturdy
cabin of gray logs chinked with white plaster, with a solid field stone
chimney and a sloping roof drifted over with pine-needles. She slid
from the saddle and stood upon the rugged doorstep. Here was her
house, her very own!
It was a larger dwelling than she had expected and very solidly
and substantially built. She found that wooden bars had been nailed
across the doors and windows, and she had, moreover, forgotten to
obtain the keys from Dan O’Leary, so that she could not go in. She
could, however, peep through the easement windows and see the
low-ceilinged rooms, the rough stairs, and the wide fireplace. The
big trees nodded overhead, the roar of the waterfall came from
beyond the house, the creek, rushing and tumbling, slid away down
the mountainside. Somebody had planted pansies on both sides of
the step, pansies that crowded and jostled each other as they only
can in the cool air of the high mountains, spreading sheets of
gleaming color over the barren soil. With a quivering sigh, Beatrice
sat down upon the step.
“Mine!” she said aloud, just to see how it would sound. “Mine!”
It would take a long time to explore the place thoroughly.
“I must be able to tell Nancy about every bit of it,” she told
herself.
Yet first she sat very quietly, for a little, on the rough stone step.
She had hurried up the hill, eager to see the new place; she had
been hurrying for the last two days, getting the house in the village
settled; she had hurried before the journey: when indeed had she
not been hurrying? It was very pleasant to sit so still and let the
silent minutes march by to the tune of rustling pine branches and
the murmuring waterfall. As she sat looking down into the valley,
time seemed very big and calm and empty, instead of bustling and
full.
 She rose at last to go on with her explorations. Behind the cabin
was the tumbling cascade that identified the place, a plunge of
foaming waters over a high ledge with a still black pool below, shot
with gleams of sunshine and full of darting trout. Beyond the stream,
almost hidden from sight by the high slope of the ravine, was the
roof of another house, a larger one than hers, with a whole group of
chimneys sending forth a curl of smoke to indicate that here were
neighbors. Looking up the course of the brook she could see where
the dense shadows of the pine grove ended and the waters ran in
brighter sunshine on the higher slope.
“I should like to see what it is like up there,” she thought, “but I
must be quick; it is getting late.”
She went scrambling up the rocky slope, feeling a little breathless,
but forgetting entirely that in such a high altitude haste is far from
wise. In a moment her lungs seemed entirely empty and her heart
began to pound against her side, but she pressed on, determined to
reach a certain high rock before she turned back. It was a rash
desire, for presently she was obliged to lie down upon the rough
grass to gasp and rest and gather herself together for another effort.
She got up to struggle forward again, for she was not used to
abandoning a fixed purpose, but after a few yards she was forced to
lie down once more, panting and completely exhausted.
“I don’t believe I understand the Rocky Mountains,” she reflected
as she lay, limp and flat, looking across the barren valley, the
sparsely wooded slopes, to the rising peaks opposite. She had been
accustomed to mountains like the Adirondacks, round and covered
thick with forest almost to the summit, friendly heights that invited
one to climb them. It was a far cry from them to the precipitate
slopes of Gray Cloud Mountain.
When she had recovered a little she gave up her project and slid
humbly down the steep way she had come. Buck, with his bridle
over the post at the cabin door, whinnied an anxious welcome as she
came back to him. He had been searching for tufts of grass between
the stones, and had also nipped at the pansies, but had found them
not to his liking. His impatience, as well as the creeping shadows in
the valley below, reminded her that evening was near despite the
clear sunshine higher up the mountainside. Reluctantly she mounted
and, with many a glance backward at her house, rode down the
trail.
Through an opening in the trees Beatrice caught a glimpse, as she
descended, of the house beyond the stream. She could even see a
man ride up to the door and a girl come running out to greet him.
Then a drop in the trail hid both house and people abruptly from her
view.
The warm sun seemed to be left completely behind as she and
Buck pressed onward with all possible haste. Something new caught
her curious attention in a moment, however, and made her stop
again. To the right of the pathway, in a little clearing among the
pines, she had spied the glow of a tiny fire.
“Who is burning brush on my land?” she questioned inwardly, with
a throb of pride at the thought of her proprietorship.
Guiding her horse among the trees, she rode a little nearer to
investigate. The blaze was kindled skilfully between two stones,
evidently by the hands of some one who knew the dangers of
careless camp-fires in a pine grove. Bending over the crackling flame
was a woman, with a yellow handkerchief covering her hair and a
green shawl slanting about her hips above a shabby skirt. A big
basket stood beside her, showing that she had been gathering
berries in the wood, while an appetizing smell rising from the fire
told of a supper of bacon and fresh trout. The smoke was in her
eyes and she was, moreover, intent on balancing the frying-pan
between the stones, so that she did not see Beatrice. For this the
girl was thankful, since, after a glance at the other’s broad, brown
face, she concluded that one ill-mannered foreigner was all she
wished to encounter that day and that she would push her
investigations no further. She turned her pony to make for the path
again, but a rolling stone, dislodged by Buck’s foot, attracted the
woman’s attention. Beatrice looked back to see that the stranger had
abandoned her cooking and was standing erect, staring intently after
them.
“At least she cannot follow,” thought the girl with some relief; then
observed, with a sinking of the heart, that the woman had turned
abruptly and was hurrying down the hill through the underbrush. It
was plain that she intended to reach the road first and intercept the
horse and rider at the bridge.
 CHAPTER II
THE DEPARTURE OF JOE LING

The yellow pony, stamping and sidling, came to an unwilling stop

before the sturdy figure that blocked the way. Beatrice began to see
that the red firelight had made the woman seem unduly terrifying
and that her face, while it was sunburnt almost to the color of
leather, was merely a square, stolid one, with keen, blue eyes and
heavy, fair hair showing under the picturesque head handkerchief.
With one hard, big hand, the stranger was feeling within her dress
and, as Beatrice came close, she held up a letter.
“I saw you in town yesterday, and you looked kind. I want you to
read my letter to me; I cannot read English myself. My name is
Christina Jensen. The letter is from my boy.”
She spoke with a strong accent that, while it was somewhat like
that of the man from whom Beatrice had asked the way, was not
unpleasant, for her voice was rich and clear. The girl thought as she
looked into the upturned face, that she had never seen such eager,
appealing eyes.
“You can’t read?” Beatrice exclaimed, forgetting politeness in her
surprise.
“My own language, Finnish, yes, but not yours. My boy, Olaf,
made me learn to talk English plain, but I was always so busy with
my two hands I could not learn to read or write. Read, read, please,
before it is too dark to see the letter.”
Beatrice spread out the paper on the pommel of the saddle.
“Why,” she said, glancing at the date, “it is nearly a year old!”
“Yes,” returned the woman nodding heavily, “ten months ago he
wrote it from his ship in Marseilles. I have nearly worn it out carrying
it around and having it read to me. But it is only kind people I ask to
read it now, for some begin to say, ‘Like father, like son; your Olaf
will never come back.’”
“Was his father a sailor too?” the girl asked.
“Yes, but he sailed away from our home in Finland when our boy
was only a month old, and I never heard from him again. It was
nearly a year later that we learned how his ship had been wrecked
on the voyage to Japan. I brought my boy to this country then
where I could support him better, and what a credit and a comfort to
me he was. He was wild to go in the navy when the war began, but
he was just too young; so it was not till last year that he slipped
away, as I had always feared he would. He hardly even said good-by
to me, and this is my only letter from him. But I talk too long; you
will not be able to see.”
Once more Beatrice turned to the paper and began:
“My dear mother: I expect you think I am never going to send you
a letter—”
She read through to the end, thinking that it sounded affectionate
but contained little news beyond the fact that the writer was going
to China.
“He gives an address to send an answer,” she observed as she
folded the letter and handed it back. “What did you write to him?”
To her surprise she saw big tears stand suddenly in Christina’s
eyes.
“Ah, Thorvik would not let me, and I couldn’t write myself,” she
said. “And my Olaf is such an American, he cannot read my
language. That is perhaps why he has not written again and has not
come home.”
Then, seeing Beatrice’s puzzled look, she explained more fully,
although it was difficult to make plain her foreign notion that women
are subject to the men in their houses.
“Thorvik is my brother, once a good Finn like myself, but now—oh,
so different. He was to come to America some years ago, but the
war broke out over here and he went, instead into the Russian army.
Now that there is peace he has come to us, but how that time had
changed him! He is full of wild talk of revolution, and tyrants and
destroying every thing. He and Olaf never agreed. It was what made
my boy unhappy at home, and, though I did what I could, Olaf went
away from us at last.”
Beatrice leaned forward in her saddle with sudden interest.
“Do you live in a little cottage half-way up the hill above Ely? That
man I saw there when I rode by—is that your brother?”
Christina nodded.
“And if you could write to your son,” the girl pursued, “what would
you say?”
“I would say, ‘Come home,’” cried Christina. “Over and over I
would say, ‘Come home. If it is only for a week or a day between
voyages,’ I would say, ‘come still, no matter what happened before
you went away.’”
Beatrice felt in the pocket of her riding skirt. There were a note-
book and pencil there, she felt sure, for she had made a list of
supplies to be bought in the village before she set out on her ride.
“Do you want me to put down the address and write to your son
for you?” she offered.
“Oh, if you would!” cried Christina. “And you would never tell
Thorvik?”
“There is no danger of that,” Beatrice assured her. “And I think
somehow that your boy will come back.”
She could not tell, herself, what made her offer such a definite
opinion.
There was something she liked about the words of the letter. “I
went ashore at Marseilles, and it is such a strange place that before
I had been there an hour I wanted to stay a year. But loafing doesn’t
suit me, so I am off again for Hong-Kong, but I’ll not forget you,
Mother, not even on the other side of the world.”
She folded the worn page once again, gave it to Christina, and
rode on. To her own surprise, she had that pleasant, satisfied
feeling, that comes with the making of a new friend. After a few
rods, she turned to look back and saw the Finnish woman still
looking after her. Beatrice raised her hand in a quick gesture of
leave-taking. It was a slight move but it had important
consequences, since it seemed to cement their regard for each other
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