Weed Biology and Climate Change 1st Edition

Lewis H. Ziska pdf download

https://ebookgate.com/product/weed-biology-and-climate-
change-1st-edition-lewis-h-ziska/

Get the full ebook with Bonus Features for a Better Reading Experience on ebookgate.com
Instant digital products (PDF, ePub, MOBI) available
Download now and explore formats that suit you...

Molecular Biology of Weed Control 1st Edition Jonathan

Gressel (Author)

https://ebookgate.com/product/molecular-biology-of-weed-control-1st-
edition-jonathan-gressel-author/

ebookgate.com

The Economics of Climate Change and the Change of Climate

in Economics 1st Edition Kevin Maréchal

https://ebookgate.com/product/the-economics-of-climate-change-and-the-
change-of-climate-in-economics-1st-edition-kevin-marechal/

ebookgate.com

Postmodern Climate Change 1st Edition Leigh Glover

https://ebookgate.com/product/postmodern-climate-change-1st-edition-
leigh-glover/

ebookgate.com

Controlling Climate Change 1st Edition Bert Metz

https://ebookgate.com/product/controlling-climate-change-1st-edition-
bert-metz/

ebookgate.com
Human Rights and Climate Change 1st Edition Stephen
Humphreys

https://ebookgate.com/product/human-rights-and-climate-change-1st-
edition-stephen-humphreys/

ebookgate.com

Informing an Effective Response to Climate Change National

Research Council 1st Edition America'S Climate Choices:
Panel On Informing Effective Decisions And Actions Related
To Climate Change
https://ebookgate.com/product/informing-an-effective-response-to-
climate-change-national-research-council-1st-edition-americas-climate-
choices-panel-on-informing-effective-decisions-and-actions-related-to-
climate-change/
ebookgate.com

Organizational Change Creating Change Through Strategic

Communication 1st Edition Laurie K Lewis

https://ebookgate.com/product/organizational-change-creating-change-
through-strategic-communication-1st-edition-laurie-k-lewis/

ebookgate.com

Climate Change and Food Security David B. Lobell

https://ebookgate.com/product/climate-change-and-food-security-david-
b-lobell/

ebookgate.com

Science vs Climate Change 1st Edition Nick Hunter

https://ebookgate.com/product/science-vs-climate-change-1st-edition-
nick-hunter/

ebookgate.com
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

Weed Biology and Climate Change

Weed Biology and Climate Change Lewis H. Ziska and Jeffrey S. Dukes
© 2011 Blackwell Publishing Ltd. ISBN: 978-0-813-81417-9
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

Weed Biology and Climate Change

LEWIS H. ZISKA, Ph.D.

Plant Physiologist
Crop Systems and Global Change Laboratory, USDA
Agricultural Research Service
Building 1, 10300 Baltimore Avenue
Beltsville, MD 20705

JEFFREY S. DUKES, Ph.D.

Associate Professor
Department of Forestry and Natural Resources, and
Department of Biological Sciences
Purdue University, 715 West State Street
West Lafayette, IN 47907

A John Wiley & Sons, Ltd., Publication

P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

Edition first published 2011

C 2011 Blackwell Publishing Ltd.

Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has
been merged with Wiley’s global Scientific, Technical, and Medical business to form Wiley-Blackwell.

Editorial Office
2121 State Avenue, Ames, Iowa 50014-8300, USA

For details of our global editorial offices, for customer services, and for information about how to apply for
permission to reuse the copyright material in this book, please see our Website at
www.wiley.com/wiley-blackwell.

Authorization to photocopy items for internal or personal use, or the internal or personal use of specific clients, is
granted by Blackwell Publishing, provided that the base fee is paid directly to the Copyright Clearance Center,
222 Rosewood Drive, Danvers, MA 01923. For those organizations that have been granted a photocopy license
by CCC, a separate system of payments has been arranged. The fee code for users of the Transactional Reporting
Service is ISBN-13: 978-0-8138-1417-9/2011.

Designations used by companies to distinguish their products are often claimed as trademarks. All brand names
and product names used in this book are trade names, service marks, trademarks or registered trademarks of their
respective owners. The publisher is not associated with any product or vendor mentioned in this book. This
publication is designed to provide accurate and authoritative information in regard to the subject matter covered.
It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional
advice or other expert assistance is required, the services of a competent professional should be sought.

Library of Congress Cataloging-in-Publication Data

Ziska, Lewis H.
Weed biology and climate change / Lewis H. Ziska, Jeffrey S. Dukes.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-8138-1417-9 (hardcover : alk. paper)
1. Weeds–Climatic factors. 2. Weeds–Control. I. Dukes, Jeffrey S. II. Title.
SB611.Z57 2011
581.6 52–dc22
2010027035

A catalog record for this book is available from the U.S. Library of Congress.

Set in 10.5/13 pt Times by Aptara

R
Inc., New Delhi, India
Printed in [country]

Disclaimer
The publisher and the author make no representations or warranties with respect to the accuracy or completeness
of the contents of this work and specifically disclaim all warranties, including without limitation warranties of
fitness for a particular purpose. No warranty may be created or extended by sales or promotional materials. The
advice and strategies contained herein may not be suitable for every situation. This work is sold with the
understanding that the publisher is not engaged in rendering legal, accounting, or other professional services. If
professional assistance is required, the services of a competent professional person should be sought. Neither the
publisher nor the author shall be liable for damages arising herefrom. The fact that an organization or Website is
referred to in this work as a citation and/or a potential source of further information does not mean that the author
or the publisher endorses the information the organization or Website may provide or recommendations it may
make. Further, readers should be aware that Internet Websites listed in this work may have changed or
disappeared between when this work was written and when it is read.

1 2011
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

This book is dedicated to

Dr. David T. Patterson and Dr. James A. Bunce.
Scholars and Gentlemen.
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

Contents

Preface ix
Chapter 1 A Brief History of Weeds and Their Impact 1
Chapter 2 Carbon Dioxide and Global Warming: The “Green” in the
Greenhouse Effect 23
Chapter 3 An Evaluation of the Impact of Rising Carbon Dioxide and
Climatic Change on Weed Biology: From the Cell to the Plant 39
Chapter 4 An Evaluation of the Impact of Rising Carbon Dioxide and
Climatic Change on Weed Biology: Competition to Community
Composition 61
Chapter 5 Weeds on the Farm: Assessing the Role of Climate Change and
CO2 on Agricultural Productivity 85
Chapter 6 Invasive Plants and Climate Change in Natural Ecosystems 107
Chapter 7 Weeds, CO2 , Climate, and Health 127
Chapter 8 Weed Management: Herbicides 143
Chapter 9 Weed Management: The Rest of the Story 163
Chapter 10 Benefits from Weeds 181
Chapter 11 Weeds in a Time of Climate 199

Appendix 219
Index 227

vii
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

Preface

Ask people for an example of a weed, and you are likely to get a variety of responses.
For most suburban commuters, dandelion, chickweed, or crabgrass and their effect on the
aesthetic quality of their lawn might be a concern. For avid gardeners, mustard, plantain, or
clover might be a consideration. Farmers may worry most about a different group of weeds,
many of which are wild relatives of the crops they grow. For a hiker or backpacker, or an
ecologist, invasive plant species such as cheatgrass or yellow starthistle would top the list.
But there are other weeds that people rarely think about. For example, the allergist works
with pollen outbreaks from unregulated ragweed proliferation. A dermatologist may deal
with contact dermatitis from poison ivy exposure. A pharmacist might study sweet Annie,
a common roadside weed in Virginia for the production of artemesinin, a new antimalarial
drug. Drug Enforcement Agency (DEA) agents could study wild poppy production and its
impact on the global supply of Heroin.
People rarely stop to think about the full impact of weeds in human society. Weeds affect
many aspects of our daily lives, from the quantity and quality of the foods we enjoy, to the
medicine we consume, the allergies we encounter, the frequency of fires in some parts of the
world, and even the diversity of species in natural ecosystems.
There are, of course, existing tomes that assess and quantify all of these aspects of weed
biology; the goal of this book is not to duplicate previous efforts. Rather, we attempt to as-
sess and synthesize recent information regarding the unprecedented increase in atmospheric
carbon dioxide and the associated change in climate, particularly temperature and precip-
itation, in regard to basic and applied aspects of weed biology. We examine responses of
weed growth and reproduction as well as weed management, invasive weeds, public health,
ecosystem functioning, and food security.
There is still much to be learned about the responses of weeds to anthropogenic increases
in CO2 and climate change. However, we are now beginning to recognize the scale and
rapid pace of environmental perturbations caused by human society, and the implications of
these changes for weed biology. Given this pace, and the interactive dynamic between weeds
and environmental disturbance, we feel there is a critical need to begin summarizing and
publicizing what is currently known in this field.
We begin with separate overviews of weed biology and climate change, and then discuss
probable responses to climatic change and rising carbon dioxide, from the genome to the

ix
P1: SFK/UKS P2: SFK/UKS QC: SFK/UKS T1: SFK
fm BLBS066-Ziska October 13, 2010 13:28 Trim: 244mm X 172mm Printer Name: Yet to Come

x PREFACE

plant community level. We hope to provide a framework for greater insight into the likely
impacts of global change on agronomic weeds, invasive plant species in natural or less
managed plant communities, and the links between weeds and public health. We discuss the
implications of global change with respect to chemical, cultural, biological, and mechanical
weed management as well as some beneficial aspects of weeds. We attempt to highlight
crucial research questions and to highlight adaptation or mitigation strategies that could
reduce potential consequences.
We stress that this is only a beginning. We are seeking new ideas and models that can help
us define, understand, and predict weed biology and likely societal impacts of unparalleled
human-induced climatic uncertainty in the twenty-first century. In doing so, we wish to cast
our academic net not only within the weed science community, but also among a wide swath
of stakeholders, including environmentalists, agronomists, and health care providers.
To that end, we are indebted to Justin Jeffryes and Shelby Allen of Wiley-Blackwell
publishing for their invitation to prepare this book and for their advice and encouragement.
We are also grateful to our own scientific mentors, Hal Mooney at Stanford University, Jim
Ehleringer at the University of Utah, Chris Field at the Carnegie Institution for Science,
Paul Epstein at Harvard, David Patterson at Duke, Jim Bunce at USDA, and Tony Hall at
University of California, Riverside, who have fostered our appreciation of nature’s wonders
and provided a healthy dash of skepticism. We are also indebted to our colleagues who gave
generously of their time in helping put this book together, Jodie Holt, Jil Swearingen, Bethany
Bradley, Dana Blumenthal, Jack Morgan, and Hilda Diaz-Soltero. Finally, we express our
profound gratitude to our families and friends, who, while supporting us, must have wondered
how grown men could be so fascinated with weeds.

Lewis H. Ziska, Ph.D. USDA-ARS

Crop Systems and Global Change Lab

Jeffrey S. Dukes, Ph.D. Associate Professor,

Purdue University
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

1 A Brief History of Weeds and Their Impact

“. . . the sun never sets on the empire of the dandelion”

Alfred W. Crosby, Ecological Imperialism

Weed Classification

Weeds were, and are, the largest single limitation on crop yield. Before the onset of chemical
control for weeds, most of the work on the farm from June through August was hoeing; a
weed control method still practiced around the world. Without mechanization, the size of a
farmer’s holding and yield was determined by how well (and how fast) a family could weed
its land. As suggested by Zimdahl (1993), more human labor may be expended on weeding
than on any other human enterprise (Figure 1.1).
Weeds affect nearly every aspect of our lives, from the appearance of our lawns to the
quality of the food we eat, even the state of our health. Weed science, a subset of botany, is a
multidisciplinary science with the goal of a systematic understanding of both weed biology,
and more importantly, weed control. Weed scientists represent a broad spectrum of scien-
tific disciplines including plant physiologists, botanists, agronomists, chemists, molecular
biologists, biochemists, and ecologists.
But what makes a plant a weed? If we called something a weed in the northeastern United
States, would it still be considered a weed in Australia? Are there common characteristics
among different weed species? In what habitats are weeds found? How do weeds spread?
What makes a weed harmful? How do we measure harm? How do we manage weeds
currently? How will we manage them in the future? In this first chapter we will explore these
basic concepts of weed biology.
A weed is a weed is a weed? One of the greatest difficulties in introducing the subject of
weed biology is a clear understanding of what is meant by “weed.” The term “weed” does
not exist in nature; rather, plant species assume the mantle of “weed” when classified as such
by human society.
Given the diversity of societies, many weed science books will list a number of cultural
and scientifically derived definitions. While varied in scope, these characterizations fall
into two broad categories: “an unwanted or undesired plant species” and “early vegetation
following a soil disturbance.” The first classification is used by many laypersons as well as
agronomists and weed scientists, the latter is a definition widely accepted by ecologists and

Weed Biology and Climate Change Lewis H. Ziska and Jeffrey S. Dukes
© 2011 Blackwell Publishing Ltd. ISBN: 978-0-813-81417-9

1
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

2 WEED BIOLOGY AND CLIMATE CHANGE

Figure 1.1 The author’s (LHZ) future in-laws working on a farm near Samantha, Alabama, during the early 1900s.
Farm work, particularly hoeing weeds, was the main activity during the summer months, and the principal reason school
children were given summers off. (Photo courtesy of Mollie Guy of Tuscaloosa, Alabama.)

environmentalists. There is also a third definition—that a weed is a plant whose virtues have
yet to be discovered—suggesting that if a use were found for a weed species, it would cease
to be a weed. Yet, many of the most pernicious weeds were introduced specifically because
of their virtues (e.g., forage crops such as Johnson grass1 and kudzu, as well as aesthetic
species such as purple loosestrife).
Because the characterization of a weed is determined by human activity, the definition of
a weed is fluid; it can vary by time and space, and certainly by culture. For example, jim-
sonweed, in addition to being a common weed of roadsides, is also considered an important
medicinal plant for native peoples of the southwestern United States. However, certain plants
are universally reviled because it is acknowledged among disparate regions and cultures that
those plants can cause widespread economic or environmental damage. One such plant is
Canada thistle, considered among the worst weeds in North America (Skinner et al., 2000)
for its ability to invade, colonize, and out-compete native and agricultural vegetation. In

1
Common names of weeds will be used throughout the book, scientific names, Bayer and WSSA codes are
referred to in the appendix
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 3

these cases, the definition of a weed is less in doubt because the impact of these plants is
universally recognized.
Ultimately, the definition of “weed” is anthrocentric and therefore flexible; however,
“weeds” in many societies share certain biological characteristics.
The following is a synthesis of some of these characteristics, most notably those presented
in Zimdahl (1993) and Radosevich et al. (1997). No single weed possesses all of these
attributes, but these characteristics encompass biological traits associated with “weediness.”

1. Long-term seed survival in the soil, including resistance to microbial degradation. In

addition, weed seeds can germinate over a wide range of environmental conditions. Soil
disturbance may be needed for germination.
2. Rapid growth following germination.
3. Mechanical (e.g., spines) and chemical (e.g., poison) barriers to herbivory during vege-
tative growth.
4. Grow over a wide range of environmental conditions, including extremes of temperature,
water, and nutrient availability; high environmental plasticity.
5. Flower early during vegetative growth. Time from flowering to seed production can be
rapid (e.g., 2 weeks from flowering to mature seed for Canada thistle).
6. While some weeds are self-compatible, most cross-fertilize between individuals, usually
using unspecialized pollen carriers or wind. As a consequence, seeds produced by any
one plant can be very genetically diverse.
7. Weeds may produce more than one seed crop per year. Individuals can produce tens of
thousands of seed in a given year.
8. Reproduction, particularly for perennial weeds, may include both sexual (floral) repro-
duction, but also asexual reproduction from roots and/or rhizomes.
9. Weeds may have highly effective seed dispersal mechanisms, from wind (dandelion) to
animals (cocklebur).

Life Cycle
Weeds can also be classified on the basis of their life history. For instance, knowing whether
a plant is an annual, biennial, or perennial can help weed scientists determine where a weed
could become a problem.
An annual is a plant that completes its entire life cycle (from seed germination to seed
development) in less than 1 year. Annuals grow quickly, and many are prolific seed producers.
Summer annuals germinate in spring, grow in summer, flower, and die with the onset of frost
in the fall. Examples of summer annual weeds include velvetleaf, pigweeds, and foxtail
grasses. Winter annuals, in contrast, germinate in fall or early winter, flowering and maturing
in spring or early summer of the following year. Examples of winter annual weeds include
chickweed, shepherdspurse, and cheatgrass.
Biennials, as the name suggests, are plant species that complete their life cycle over a 2-
year period. These species typically form a vegetative rosette during the first year of growth,
become quiescent over winter, and then bolt during the following spring, forming a tall
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

4 WEED BIOLOGY AND CLIMATE CHANGE

Figure 1.2 Chickweed, curly dock, and field bindweed as common examples of annual, biennial, and perennial weeds.

inflorescence with prodigious seed development. Examples of biennials include curly dock,
garlic mustard, bull thistle, and musk thistle.
Among the most damaging of weeds are perennials, those that are long-lived, multi-year
plant species. These can be divided into two subgroups: simple and creeping (Zimdahl,
1993). Simple perennials reproduce and spread primarily by seed, whereas creeping peren-
nials can spread by both sexual (i.e., seeds) and asexual (i.e., rhizomes and roots) means.
Examples of simple perennials would include common lawn weeds such as dandelion or
plantain, or weeds of wet damp locations such as curly dock. Creeping perennials include
weeds associated with contact dermatitis such as poison ivy (rhizomes and stems), and
common weeds in pastures such as leafy spurge (rhizomes) and Canada thistle (roots)
(Figure 1.2).

Classification by Physiology
Differences in photosynthetic pathway provide another means to classify weeds. This type
of classification will also be important when we examine ongoing increases in atmospheric
carbon dioxide associated with climate change in later chapters.
Initial investigations of how plants acquire carbon led to the discovery that one of the
first products of photosynthesis was the production of phosphoglyceric acid, a three-carbon
compound (Calvin, 1949). For the vast majority (approximately 95%) of plant species it
was discovered that this type of photosynthesis (termed C3 because the first stable product
had three carbons) was the sole means by which carbohydrates were synthesized from
light, energy and water. However, in the 1960s, researchers discovered other photosynthetic
pathways, most notably the C4 pathway, where the first products of photosynthesis are
four-carbon sugars such as oxaloacetate, malate, and aspartate. Although these C4 plants,
principally tropical grasses, constitute only about 4% of all plant species, a higher percentage
of this type of photosynthesis appears to occur among the world’s worst weeds (e.g., Holm
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 5

et al., 1977). A third type of photosynthetic pathway, crassulean acid metabolism or CAM,
occurs in less than 1% of plant species. While succulent species use CAM photosynthesis,
major weeds do not.

Classification by Habitat
One of the most well-recognized habitats for weeds is cropland. However, because crops
cover a wide range of different environments, there may not be a specific set of characteristics
associated with crop weeds per se. Interestingly, the most competitive weeds in a given
cropping situation are often simply “imitators” or wild relatives of the crop. They are selected
for by cultural practices such as planting date, fertilizer application, selective herbicide
usage, etc. which while favoring the crop, also select for those weeds that are closely
related physiologically, morphologically, or phenologically. Examples of crops and their
weed relatives include potatoes and nightshade (both in the genus Solanum), rice and wild
or red rice, sorghum and shattercane (both Sorghum bicolor), oat and wild oat.
Rangelands constitute those land areas characterized by grassland and shrubland used
(although not exclusively) for animal grazing. Because of their use, classifications in range-
lands are associated with those weeds whose presence is either directly (e.g., poisonous) or
indirectly (competes with desired grasses) harmful to large animal (sheep and cattle) grazing.
Examples of typical rangeland species include leafy spurge and yellow starthistle.
Forests have another unique set of weed species associated with them. James Miller of
the U.S. Forest Service has identified over 33 plants or groups that are spreading rapidly
through southern forests (Miller, 2003). As might be expected, vines that compete directly for
sunlight such as kudzu, Oriental bittersweet, or English ivy are considered among the worst
nonnative vines, but other perennial weeds such as Chinese and Japanese privet, nonnative
roses, and sacred bamboo are also among the worst forest weeds. In addition, certain tree
species can also be considered weeds and among these, nonnative trees such as Princess tree
and tree of heaven are considered common weeds in forest environments.
Riparian weeds are those species that are found in wet, poor drainage areas. They are
adapted to anaerobic soils and may rely on water as a means to spread by seed or asexually.
Riparian weeds include purple loosestrife, curly dock, giant reed, or salt cedar.
Aquatic weeds are those weeds adapted to living directly in the water. They can be floating,
submersed, or immersed (submersed but with a root system). They can reduce water quality
and water supply as well as pose problems for navigation or power generation. Some of the
most prevalent aquatic weeds include water hyacinth and hydrilla (Figure 1.3).

A Few Basic Principles of Weed Ecology

Weed ecology is that aspect of weed biology that studies weed distribution, growth, devel-
opment, reproduction, and population dynamics of weed species in managed (human) and
unmanaged (natural) plant communities.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

6 WEED BIOLOGY AND CLIMATE CHANGE

1 2 3 4 5
Figure 1.3 Weeds by habitat: (1) common lambsquarters in a soybean field; (2) Canada thistle as a rangeland weed
in South Dakota; (3) tree of Heaven, a common weed in urban areas; (4) purple loosestrife, a common riparian weed in
marshes; (5) water hyacinth, a common aquatic weed. (Photos are all from USDA-ARS Image Gallery.)

Seed Biology
We can begin with seed, and an appreciation of the ability of weeds to produce them. The
number of weed seeds present in soil can be enormous. Koch (1969) has derived figures that
indicate between 30,000 and 350,000 weed species per square meter (or 300 million to 3.5
billion seeds per hectare). An estimate of the number of seeds produced for a single weed
can be in the hundreds of thousands (Table 1.1).
What is the fate of seed once it enters the soil? Harper (1977) envisioned the soil as a seed
“bank” in which “deposits” or “withdrawals” are made. An example of a deposit would be
seed rain, whereas withdrawals (seed removal) would occur by germination, deterioration, or
predation. The concept of a seed bank is a very useful one for weed ecology since it provides
key information regarding seed diversity and longevity as well as long-term information as
to efficacy of weed management.
Seed longevity in turn, will depend on a number of factors and their interaction. Obvi-
ously, how much seed a given species produces is an important consideration as is soil type.
However, seed decomposition, predation, and dormancy/germination are also key consider-
ations. Overall, observations indicate that while seeds from grass and crop species are rarely
long-lasting within the seed bank, weed seeds may last for hundreds of years if not millennia
(Odum, 1965).Weed seeds are almost always present in the soil.
Seed germination and emergence is dependent on dormancy. Although there is a recog-
nized physiological aspect of dormancy (seed ripening), most dormancy is imposed by the
environment. Dormancy, in turn, can be broken by a given environmental factor or com-
bination of factors, usually those that occur in seasonal cycles. For example, freeze/thaw
combinations may be necessary to remove seed coat restrictions. Alternatively, many weed
seeds need light (Sauer and Struik, 1964), which is usually abundant following a physical
disturbance in the environment.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 7

Table 1.1 Seed production (per plant) and qualification (+ or −) as to the presence of asexual
reproduction for selected weed species.
Seed Asexual Comments

Canada thistle (Cirsium arvense 530–5,300 ++ Extensive below ground root

[L.] Scop.) system 3 m horizontal and
vertical lengths
Common lambsquarters 9,000–370,000 − Prolific seed producer
(Chenopodium album L.)
Common mullein 200,000–300,000 − Prolific seed producer
(Verbascum thapsus L.)
Common ragweed 3,000–5,000 − Agronomic weed and public
(Ambrosia artemisiifolia L.) health issue
Dalmation toadflax (Linaria up to 500,000 ++ Extensive deep root system
dalmatica L. [Mill])
Diffuse knapweed (Centaurea up to 18,000 − Not asexual, but can regrow from
diffusa) cut crown
Field bindweed (Convolvulus 25–300 ++ Extensive rhizome growth and
arvensis L.) establishment of large colonies
Musk thistle (Carduus nutans L.) up to 100,000 − Normally biennial, can be annual
with warmer temperatures
Perennial sowthistle (Sonchus up to 13,000 ++ Can spread extensively by roots
arvensis L.)
Purple loosestrife (Lythrum up to 3,000,000 − Can regrow from woody
salicaria L.) rootstock if shoots are cut
Quackgrass (Elymus repens L.) up to 8,000 ++ Extensive rhizome production,
can form large colonies
Wild oat (Avena fatua L.) 200–700 − Common weed in cereal
production
Yellow starthistle (Centaurea up to 150,000 − Can regrow if shoots cut
solstitialis L.)

Reference : www.invasive.org.

Seed germination is associated with rapid metabolic activity, radicle emergence, and
then shoot appearance. Germination is a perilous ecological proposition, and natural selec-
tion has caused the timing of germination to coincide with favorable environmental condi-
tions (Probert, 1992). However, even with such selection, given the large seed populations
of some weed species, it is evident that many of them fail to germinate, decompose, or
get eaten.
What then are “favorable environmental conditions” that promote weed seed germination
within the seed bank? Whether human or natural, any disturbance exposes soil to light,
alters water and temperature regimes, and, assuming temperatures above thermal limits, can
allow seeds to break dormancy and germinate. As we have seen, weeds can be a primary
constituent of the seed bank; such physical (abiotic) disturbances are therefore a key factor
in their establishment within the plant community.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

8 WEED BIOLOGY AND CLIMATE CHANGE

Emergence and Competition

Given physical disturbance of the soil as a signal for extensive seed germination, what factors
following emergence from the soil are associated with weed species success? What, in other
words, makes some weeds more competitive than others?
It is difficult to provide a complete answer to these questions, and scientists are still
addressing many aspects of competition. However, in general, success at competition reflects
the ability of a given species to acquire a limited set of resources at a greater rate than its
neighbors, whether those neighbors are a different species (interspecific competition) or the
same species (intraspecific competition). How these resources are acquired depends on both
genetic hard-wiring (e.g., rate of leaf development, nitrogen use efficiency, etc.) and spatial
and temporal variability of physical resources (e.g., nitrogen distribution in the soil, rainfall,
temperature, etc.). In general, plant competition is for abiotic resources, principally light,
water, and nutrients between or within species.
Because light varies in duration, intensity, and quality, and must be used immediately or lost
permanently, those plants that intercept light and shade their neighbors are at a competitive
advantage. Plants invest heavily into new leaf formation, particularly after emergence. Indeed,
a simple comparison of leaf area ratio among seedlings can be used as one measure of
early competitive success between weeds and crops (Kropff and Spitters, 1991). However,
too much investment in leaves at the expense of stems or branches may also result in a
disadvantage, since height is an important consideration in light interception. Overall, in
many cropping situations where water and nutrients may be nonlimiting, light competition
may be one of the principal factors influencing weed–crop interactions (Donald, 1961) and
reducing crop productivity (e.g., oat and wild oat; Cudney et al., 1991).
Water availability varies greatly by region, and competition for water among seedlings
can be severe in areas where natural rainfall is low or inconsistent (e.g., rangelands and
savannah). Less competition is usually associated with irrigated systems, although variation
in delivery times and water quality (e.g., salinity) may be an additional feature related to
competitive success. Overall, there are three factors associated with competition for water
as suggested by Radosevich et al. (1997): (a) variation in the total amount and temporal
availability of water inputs into the soil; (b) soil volume available to the plant, i.e., the rate
of root development and exploration of that volume; and (c) physiological factors related
to the exchange between carbon acquisition and water loss. For some weedy species, rapid
depletion of available water may be an effective strategy for competition with crops (e.g.,
cocklebur and soybean; Scott and Geddes, 1979); alternatively, a number of common weedy
species have the C4 photosynthetic pathway, which increases overall water use efficiency
(the ratio of CO2 uptake to H2 O loss) and may be at an advantage within the plant community
if water becomes limiting. In general, success in competition for water will be dependent
on the relative abilities of the individual species to obtain the resource and to tolerate water
deficits (Table 1.1; Patterson, 1995).
Nutrients are a recognized environmental resource needed for plants to complete their
life cycle. Nutrients are divided into two broad categories based on the relative needs
of the plant, namely, macronutrients (those needed in large amounts), such as carbon,
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 9

oxygen, hydrogen, nitrogen, potassium, calcium, phosphorous, magnesium, and sulfur; and
micronutrients (needed in lesser amounts) such as iron, chlorine, copper, manganese, zinc,
molybdenum, and boron. Competition for these nutrients is both dynamic and complex since
nutrient availability is dependent on soil characteristics, most notably pH, and aspects of
plant morphology such as root development, that are species specific (Patterson, 1995). It
is difficult then to predict a precise competitive outcome for an explicit nutrient deficiency.
Nevertheless, there are some generalizations that can be made. First, it is clear that weeds
utilize nutrient resources at least as rapidly as many crop species. This suggests that weed
removal should result in greater nutrient availability to the crop species. Second, there is
evidence that weeds may be more efficient at removing nutrients from the soil than cultivated
plant species (Vengris et al., 1955).
The previous examples are meant to be illustrative, and are by no means a complete
synthesis of what is known regarding competition. For example, while nutrients, water,
and light are presented as individual physical limitations, it is common for one or more
to be limiting simultaneously. In addition, most competition experiments have focused on
suboptimal conditions, not on circumstances where such factors may be in excess (e.g.,
flooding and agronomic fertilization). Yet, differential responses to an overabundance of
resources may also have significant implications for plant–plant interactions.

Weed Reproduction and Spread

Having examined how weeds survive in the seed bank, germinate, emerge, and compete suc-
cessfully, we can turn our attention to reproduction and distribution within the environment.
How do weeds become distributed in the environment?
One reproductive method, common among many perennial weeds, is through vegetative
or asexual reproduction. Vegetative portions of the plant capable of regeneration include
stolons, long, slender stems that grow parallel to the soil surface (e.g., Bermuda grass);
rhizomes, stems that grow underground, while producing adventitious roots and shoots
(e.g., quackgrass and poison ivy); tubers, which are extensive belowground storage organs
(enlarged rhizomes) that possess axillary buds (e.g., Jerusalem artichoke); bulbs, modified
underground buds capable of regeneration (e.g., wild garlic and wild onion); corms, enlarged
underground perennial stems for carbohydrate storage (e.g., wild onion); and roots (usually
horizontal that can give rise to separate shoots (e.g., Canada thistle and field bindweed).
Although many perennials do produce vegetatively, most, if not all, also reproduce sexually
through flowering. Hence, such weeds can maintain both a high density of individuals for
a given surface area, as well as spread via seed dispersal. It is important to distinguish the
seed bank and the presence of vegetative bud reserves since both are an aspect related to
establishment, dominance, and competition from the soil. In general, buds are representative
clones from a single plant, whereas the population of buried seeds represents a broader range
of genotypes. Consequently, vegetative reproduction should be at an advantage when envi-
ronmental conditions are stable, whereas seed genotypes would be favored with continuous
disturbance. This would suggest that after a soil disturbance, seed genotypes would dominate
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

10 WEED BIOLOGY AND CLIMATE CHANGE

initially, followed by the appearance of herbaceous perennial weeds with high vegetative bud
reserves. Such vegetative reserves can be impressive. For example, within 2 years, Canada
thistle plants can produce over 66 feet (20 m) of new roots (Parsons, 1992).

Getting the Seed Out

As we have already observed, one of the characteristics of weed species is prodigious seed
production. However, widespread distribution of these seeds, through a wide array of vectors,
aids in their spread and negatively impacts their control.
A number of weed species have plumed seed for wind dispersal. These include dandelion,
Canada thistle, mare’s tail, and a number of weeds within Asteraceae. The plume, or pappus,
is the “parachute” used for wind dispersal, with the extent of wind displacement related to the
ratio of seed weight to pappus size. That is, those species that produce a low ratio of achene
diameter to pappus size have higher “hang-time” and can disperse farther from the mother
plant. Plant size, particularly the height of seed release, is also positively correlated with area
of dispersal. Tall plants (those over 1 m in height) can distribute seed as far as 5 km (Dauer
et al., 2007) depending on wind velocity and degree of infestation. In addition to disseminat-
ing seed directly, wind can also move senescent or detached portions of a plant, with subse-
quent distribution of seed over a wide area. For example, tumbleweeds such as Russian thistle
can travel several kilometers while dispersing 60,000 seeds per plant (Stallings et al., 1995).
A number of weed seeds without any special morphological modifications can travel by
water. This is a particular concern in agricultural areas where irrigation is common. An
early study by Eddington and Robbins (1920) found a total of 81 different weed species in
irrigation ditches in Colorado. A study done by Wilson (1980) found a total of 77 different
plant species in three main irrigation canals over two seasons, collecting a total of 30,346
seeds. Interestingly, many weed seeds are able to survive long periods of time submerged in
water. For example, seed of field bind weed had 55% germination, even after 54 months under
water (see Table 5.3; Zimdahl, 1993). Water then, can be an effective means to distribute
weed species over a large geographic area.
Many weed seeds are characterized by having various hooks and/or barbs on the seed
coat (e.g., puncturevine and cocklebur). Others may possess a sticky coating or resin (e.g.,
gumweed). These features are commonly associated with mechanical dispersal of weed seeds
by animals. Spread of some weed species in the western United States, for example, can be
traced to grazing areas of sheep and cattle (Crafts, 1975). In addition to such mechanical
dispersal, animals may also distribute seed directly by ingestion followed by partial digestion.
After passing through the alimentary canal, seed can then be spread via feces. Harper (1977)
and King (1966) provide a number of examples of seed dispersal by birds, rodents, and wild
and domesticated ruminants. Bird distribution in particular can move seeds over thousands
of miles (Proctor, 1968).
Human activities, particularly in agriculture, are a well-recognized vector in weed seed
dispersal. Farm machinery, such as plows, harrows, and even tires can spread weed seed
well beyond its original source. Farming practices influence which weed species become
established. For example, in rice farming, direct seeded rice is associated with the spread
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 11

of wild or “red” rice because the seed of red rice is often indistinguishable from that of the
cultivar. Conversely, rice that is transplanted into flooded conditions is associated with weeds
that are tolerant of such wet or paddy environments such as Echinochloa species.
In addition to farming practices, the globalization of trade has also resulted in the intro-
duction of numerous “exotic” or “invasive” weed species. For example, Japanese stiltgrass,
a forest understory weed, was introduced as packing material into Tennessee in the early
twentieth century (Fairbrothers and Gray, 1972); cheatgrass may have been introduced as a
contaminant with crop seed (Novak and Mack, 2001). Introduction of other weedy species
has been deliberate because their initial introduction was thought to be of economic benefit.
Perhaps the most notorious example is kudzu, a leguminous perennial vine; now spread over
8 million acres in the southeastern United States, whose properties as forage and feed for
cattle attracted the attention of the USDA, which heavily promoted the planting of kudzu as
late as the 1940s.

The Harm that Weeds Do

So far we have discussed a number of principles that can be used to characterize weeds and
their ecology. This may seem like a pointless exercise, since many people do not consider
plants to be “bad.” However, it is important to acknowledge the harmful nature of weeds and
to recognize that “green” is not always good.

Food
Agriculture is the principal provider of food for the human population. Of the 250,000+ plant
species on Earth, only a few species have been able to be specifically cultivated for human
consumption, and of those, three cereals, rice, wheat, and corn supply the global population
with approximately 50% of their calories. Overall, more than 90% of the population is
supplied by less than 20 plant species (Diamond, 1997). Although one can argue the benefits
of modern agriculture (ability to feed large numbers with small amounts of land) versus its
detriments (large inputs of fertilizer and water contaminants), there is no question that the
current population of the world (6.9 billion) is very much dependent on modern agricultural
techniques and the maintenance of high crop yields.
Any impediment on these yields therefore has enormous implications for the ability of a
region or country to provide sufficient food for its population. The three major biological
barriers to maximal crop yields are insect pests, weeds, and plant pathogens. Among these,
weeds are the largest limitation to maintaining crop yields. There are many times more
weed species than crops; for example, in the United States, there are approximately 10 weed
species that compete with each crop species (Bridges, 1992).
Weed/crop competition is a major factor limiting food production worldwide. For example,
in China 10 million metric tons of rice is estimated to be lost annually due to weed competition
(Zhang, 2001), enough rice to feed 56 million people for 1 year (Labrada, 2007).
In addition to their direct effects on reducing crop yields, some weeds host damaging
insects or pathogens. Tall morning glory, for example, is a host for leaf blight, cocklebur for
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

12 WEED BIOLOGY AND CLIMATE CHANGE

stem rust; insects such as the corn borer may use Russian thistle as a means to spread curly
top. Other weeds may be hosts for different nematodes that damage soybean and corn.
While negatively impacting yields, weeds may also affect quality. The presence of wild
oat or red (weedy) rice can reduce milling quality of rice and wheat flour, and can reduce
brewing quality for beer. The presence of weeds within forage crops such as hay can also
reduce both the protein concentration and the economic return for growers.
Negative effects of weeds on pasture and forage crops can also reduce animal quality.
In western rangelands, some widespread weed species such as larkspur, leafy spurge, and
yellow starthistle are poisonous or do direct physical damage to animal grazers. Such damage
is considerable. For example, it has been estimated that leafy spurge reduces the livestock
carrying capacity of pasture and rangeland by 20–50% with a subsequent loss of $35–45
million in U.S. beef and hay production (Bangsund and Leistritz, 1991).

Water
Many aquatic weeds complicate water utilization and management. Pervasive water weeds
such as hydrilla (water thyme) in the southeastern United States and water hyacinth and com-
mon coontail globally can form large mats that interfere with transport of irrigation water
and cooling water for hydroelectric plants. Such weeds may also interfere with recreational
activities such as boating or fishing. Riparian weeds may also be heavy users of water, with a
subsequent strain on water availability (Anderson, 1982). For example, salt cedar invasions
in riparian areas of the southwestern United States are often blamed for depleting fresh-
water resources needed for irrigation or urban centers (although there is some controversy
surrounding this point; Shafroth et al., 2005; Figure 1.4).

Environmental Damage
Some weeds may colonize a given habitat so successfully that native species may be elimi-
nated altogether, with a subsequent decline in species diversity. Such weeds, usually nonna-
tive for a given system, have proliferated to such a large extent that they are recognized by
a separate classification; invasive or exotic weeds. Their introduction, commonly by human
transport, results in widespread economic or environmental damage (Mooney and Hobbs,
2000). An example of the type of environmental damage related to the introduction of these
invasive weeds is alteration of the fire cycle within native plant communities. These cycles
are, in turn, accelerated by the introduction of fire-tolerant grass species such as cheatgrass,
that can produce combustive vegetative with subsequent increases in the frequency, spread,
and intensity of natural- or human-induced fire (Brooks et al., 2004). The conversion of
shrub-dominated steppes of the western United States to a cheatgrass-dominated landscape
during the twentieth century is a quintessential example of how invasive species introduction
can significantly alter plant community composition over a large geographic area.
Cheatgrass is not the only species having dramatic effects on the landscape (Dukes and
Mooney, 2004). Millions of acres of productive rangelands, forests, and riparian areas have
been overrun by weedy invaders, with a subsequent loss of native flora. The proliferation
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 13

Figure 1.4 Salt cedar as a threat to water flow in the western United States. Accumulation of salt by this species may
also alter soil salinity with negative effects on native plants. (John M. Randall, The Nature Conservancy.)

of aquatic weeds, in turn, can lead to greater eutrophication and significant disruptions in
marine biology. It has been estimated that more than 200 million acres of natural habitats
(primarily in the western United States) have already been lost to invasive, noxious weeds,
with an ongoing loss of 3,000 acres a day (Westbrooks, 1998). The invasive plant species
that are most harmful to native biodiversity are those that significantly change ecosystem
processes (e.g., cheatgrass) to the detriment of native species.

Public Health
While the interaction of weeds and public health may seem somewhat obscure, weeds can
directly affect public health through allergenic reactions, skin irritations, mechanical injury,
and toxicology. Current estimates suggest there are approximately 30 million plant-based
allergy sufferers in the United States. While many of these people are allergic to pollen of
ornamental trees and grasses, a common weed, ragweed (Ambrosia spp.) is the principal
source of plant-based allergens in the fall. A different kind of allergy, contact dermatitis or
a skin-based allergic reaction, is also associated with weeds. Some examples include the
milky sap in spurges or giant hogweed, stinging nettles, or urushiol contact with the poison
ivy group. In the latter case, it is estimated that over 2 million people in the United States
suffer annually from casual contact with poison ivy, poison oak, or poison sumac (Mohan
et al., 2006).
Mechanical injury associated with weeds, from unwittingly removing a Canada thistle
plant by hand to stepping barefoot on puncturevine seeds, can be a painful, although nonfatal
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

14 WEED BIOLOGY AND CLIMATE CHANGE

Poison oak Poison sumac Poison ivy

Figure 1.5 Poison ivy, a common weed of disturbed sites and contact dermatitis from poison ivy. (From USDA NRCS
plants database.)

experience. Ingestion of poisonous weeds, however, can result in serious illness or death.
There are over 700 plant species that are known to induce illness in humans or livestock.
Some weed species such as poison hemlock, nightshade, or castor bean are so poisonous that
even small quantities can be lethal. For example ricin, the poison contained within castor
bean, has been linked to terrorist threats, in part, because ricin has a greater potency than
cyanide (Figure 1.5).
There are other, more indirect means by which weeds impact human suffering. If weeds
are unwanted plants, then plant-based sources of narcotics, from coca to poppy should be
considered a significant threat to public health. Weeds may also indirectly alter air quality.
Kudzu, for example, emits isoprene, a chemical precursor leading to tropospheric ozone
formation (Sharkey and Loreto, 1993). Weeds can also affect the prevalence of human
disease by providing food sources for disease vectors. Pollen, for example, can serve as a
food source for mosquito larvae (Ye-Ebiyo et al., 2000), with potential interactions between
mosquito populations and weed pollen.

Aesthetics
Aquatic weeds do not just interfere with water delivery, but detract from our enjoyment
of the pleasures of nature, such as swimming in a clear lake. Avid gardeners spend many
hours removing dandelions, chickweed, carpetweed, and garlic mustard because those plants
take away from the colorful design the gardener wishes to create. Lawn enthusiasts also
spend many hours (and many dollars) in cultivating a uniform green sward. And, as any
home-owner can tell you, weeds can reduce property values. Yet, how can one quantify these
impacts? Aesthetical deterioration of the environment is universally recognized, but remains
ill-defined.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 15

1 2 3 4
Figure 1.6 Examples of weed control: (1) Dr. John Teasdale, USDA-ARS weed scientist examines hairy vetch as
potential cover crop in cultural weed control; (2) mechanical weed control through conventional tillage in soybean; (3)
use of herbicides in no till field crop; (4) Dr. Rick Bennett, USDA-ARS Plant Pathologist, examines fungi that could be
used for biological weed control. (Photos are USDA-ARS Image Gallery.)

Weed Management

Given the damage that weeds do to human activities, to the environment, and to public health,
weed management remains, even today, the principal focus of weed science. Management,
in turn, can be defined as those practices used to restrict the growth and spread of weeds. In
general, these practices focus on three main areas: prevention, eradication, and control.
Prevention refers to those actions taken to inhibit the introduction and/or establishment of
weed species into new areas. Eradication is defined as a set of measures taken to completely
remove a weed species from an area. However, complete eradication is rarely achieved, in
part, due to the high cost and effort involved in eliminating a weed species, particularly if
that species has spread over a wide (>10 ha) area. Control is the more achievable outcome
once a weed has become established, and refers to those methodologies that can reduce (but
not eliminate) weed impacts to manageable levels. There are four recognized methods of
weed control: (1) cultural, (2) mechanical, (3) chemical, and (4) biological (Figure 1.6).

Cultural
Cultural control recognizes and optimizes those practices common to good land and water
management. At the farm level, these may include using weed-free crop seed, crop rotation to
reduce the accumulation of high weed populations of certain species, and the use of smother
crops or living mulches such as hairy vetch to control weed seed emergence (Teasdale and
Mohler, 1993).

Mechanical
Mechanical control reflects some of the oldest known means to eliminate weed populations,
including hand-pulling and hoeing. However, mechanical may also refer to flooding, a typical
means of weed control in rice cultivation since rice can survive flooded conditions and many
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

16 WEED BIOLOGY AND CLIMATE CHANGE

weeds cannot; the use of nonliving material to mulch the soil, usually as a means to prevent
light interception from emergent weed seedlings; flaming or burning, utilizing controlled fire
to burn young succulent weeds; and machine tillage, such as the use of chisel plows, weed
knives, disc harrows, or some other tractor implement that breaks the soil and buries the
weed.

Chemical
By far in the United States, and in most developing countries, chemical control of weeds
through the selective application of herbicides is the most utilized practice. If applied prop-
erly and with respect to their utilization and safety, herbicides have been the most effective
means to control weeds and to limit their negative impacts, particularly in crop production
systems. For example, it has been estimated that in the United States, under best man-
agement practices, production losses associated with weeds averaged between 7 and 10%;
however, without herbicides, those losses were between 35 and 38% (Bunce and Ziska, 2000;
Table 1.2).
In 2002, approximately 78,000,000 ha of land in the United States was chemically treated
to control weeds, grass, or brush; for comparison, 34,000,000 ha were treated chemically for
disease, nematode, and insect control combined, for that same year (National Agricultural
Statistical Service, www.nass.gov; Figure 1.7).
Because of their importance, many weed science texts are written specifically to address
herbicide chemistry, focusing on physiological responses and a systematic evaluation of
application methods that can increase uptake and overall efficacy of the herbicide. Such
detail, however, is beyond the scope of this book, and we refer interested readers to excellent
books by Duke (1985) and Monaco et al. (2002).

Table 1.2 Estimated production losses due to weeds for selected crops in the United States.
Best management practices Best management practices
Crop (‘000 Mt) without herbicides (‘000 Mt)

Barley 261 (5) 1,045 (20)

Corn 19,967 (6) 99,837 (30)
Cotton 349 (8) 2,139 (49)
Potato 1,219 (6) 5,890 (29)
Rice 628 (7) 4,847 (54)
Sorghum 1,033 (8) 4,519 (35)
Soybean 5,838 (8) 27,731 (38)
Sugarcane 2,452 (9) 10.081 (37)
Wheat 3,382 (6) 11,273 (20)
Total 35,129 167,362

Production loss percentages (in parentheses) were calculated from data in Bridges (1992) and applied to
2007 production figures. Average crop losses due to weeds were 7% for best management practices and
35% for best management practices but without herbicide. Production values for sugarcane refer to bulk
cane production.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 17

2002 estimated annual agricultural use

Average annual use of

active ingredient
(pounds per square mile of agricultural
land in county) Total Percent
Crops pounds applied national use
No estimated use
Soybeans 70,699,653 69.49
0.001–0.499 Corn 7,491,973 7.36
Cotton 7,063,492 6.94
0.5–3.445 Wheat for grain 3,509,151 3.45
3.446–14.669 Cropland in summer fallow 2,514,196 2.47
Citrus fruit 2,186,741 2.15
14.67–47.984 Sorghum 1,596,879 1.57
Rice 1,057,534 1.04

Figure 1.7 Estimated annual usage of glyphosate, the most frequently used herbicide in the United States. (From 2002,
USGS pesticide use maps.)

In brief, herbicides are phytotoxic, usually organic compounds, divided into chemical
families which include aliphatic and aromatic carboxylic herbicides, the substituted urea,
carbamate, dintirophenol, dinitroaniline, s-triazine groups as well as the phosphonate/organic
phosphorous compounds such as glyphosate. Nonorganic herbicides may include salts (e.g.,
copper sulfate) as well as acetic acid. Herbicides can be either selective or nonselective
regarding the type of plant targeted, and may or may not travel to the site of action (i.e., sys-
temic). Herbicides are either applied directly to emerged plants (postemergence) or directly
to the soil (preplant or preemergence). Application of the herbicide may be as a spray in
water or oil, or as dry granules. Added surfactants such as soap often increase absorption of
the active ingredient by the plant.

Biological
Biological control is a strategy to employ a specific organism (parasite, predators, and
pathogen) that once introduced will provide a measure of weed control, usually specific to the
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

18 WEED BIOLOGY AND CLIMATE CHANGE

weed species. Biological control, if successful, can result in permanent weed control because
the controlling organism is self-perpetuating. Another advantage is that the controlling
organism, if benign, does not contribute to environmental pollution.
There are, however, a number of cautionary hurdles that must be overcome if biological
control is to be effective. If the weed to be controlled is unwanted in one area, but valued in
another, and both areas are within the same geographical region, biological control should
not be used. Introduction of the biocontrol agent must be specific to the organism to be
controlled and introduction of the controlling organism should not pose an additional envi-
ronmental threat (e.g., cane toads introduced to Australia for biological control have become
an enormous problem in themselves). The organism must tolerate similar environmental
conditions as the weed species it is directed against, so it can survive in the same range.
Overall, to produce an effective biocontrol measure requires money, time, and much effort
on the part of the research community.

Integrated Pest Management

Integrated pest management (IPM) is not a single control measure per se, but an amalgamation
of available means to monitor and control all pests (weeds, diseases, and insects) (Flint and
van den Bosch, 1981). The driving force behind IPM was to develop a synergistic approach
to monitoring and controlling pest populations. Overall, IPM is organized around six basic
tenets:

1. Determination of acceptable pest levels, with the focus on control rather than eradication.
2. Use of cultural practices as previously described, in order to maintain healthy plants
selected for a specific growing environment.
3. Monitoring of background pest levels. Regular observation is a fundamental principle of
IPM. Visual inspection, traps, and other observational methods are employed to establish
a history of potential pests for the region.
4. Mechanical control as a first line of defense. If a pest(s) reach unacceptable levels,
mechanical options such as hand-pulling or tillage are utilized.
5. Chemical control as required. Emphasis on the use of chemicals as necessary and
application of these chemicals at the most optimal time for successful weed control
(e.g., greater efficacy of herbicides applied when weeds are in the seedling stage). The
use of organic products in the field with limited environmental carryover whenever
possible.
6. Development of long-term nonchemical solutions such as biological control.

Summary

It is impossible to summarize all that is known regarding weeds in a single chapter. It is hoped
that the themes introduced here will provide an overview of the key aspects of weed biology,
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 19

particularly how they are defined, characterized, and classified; the basic principles of weed
ecology, focusing on the steps from seed germination to seed production; an overview of the
negative impacts weeds have, both in human society and in nature; and, lastly but perhaps
most importantly, methods of weed control by which these impacts can be minimized. In the
following chapters, we will explore these themes in more detail, particularly in the context
of anthropogenic climate change, the subject of our next chapter.

References

Anderson JE. 1982. Factors controlling transpiration and photosynthesis in Tamarix chinensis Lour. Ecology
63:48–56.
Bangsund DA, Leistritz FL. 1991. Economic Impacts of Leafy Spurge on Grazing Lands in the Northern
Great Plains. Agricultural Economic Report No. 275-S, North Dakota State University, Department of
Agricultural Economics, Fargo, ND.
Bridges DC. 1992. Crop Losses Due to Weeds in the United States. Weed Science Society of America,
Champaign, IL, 403 pp.
Brooks ML, D’Antonio CM, Richardson DM, Grace JB, Keeley JE, DiTomaso JM, Hobbs RJ, Pellant M,
Pyke D. 2004. Effects of invasive alien plants on fire regimes. Bioscience 54:677–688.
Bunce JA, Ziska LH. 2000. Crop ecosystem responses to climatic change: Crop/weed interactions. In: Reddy
KR, Hodges HF (eds) Climate Change and Global Crop Productivity. CABI Publishing, New York,
pp. 333–348.
Calvin M. 1949. The path of carbon in photosynthesis. VI. J. Chem. Educ. 26:639–657.
Crafts AS. 1975. Modern Weed Control. University of California Press, Berkeley, CA, 440 pp.
Cudney DW, Jordan LS, Hall AE. 1991. Effect of wild oat (Avena fatua) infestations on light interception
and growth rate of wheat (Triticum aestivum). Weed Sci. 39:175–179.
Dauer J, Mortensen DA, Van Gessel M. 2007. Spatial and temporal dynamics governing long distance
dispersal of Conyza canadensis. J. Appl. Ecol. 44:105–114.
Diamond JM. 1997. Guns Germs and Steel: The Fate of Human Societies. WW Norton and Company, New
York, 480 pp.
Donald CM. 1961. Competition for light in crops and pastures. In: Mechanisms in Biological Competition,
Soc. Exp. Biol. Symposia XV. Academic Press, New York, pp. 282–313.
Duke SO. 1985. Weed Physiology. Vol.2: Herbicide Physiology. CRC Press, Boca Raton, FL,
283 pp.
Dukes JS, Mooney HA. 2004. Disruption of ecosystem processes in western North America by invasive
species. Rev. Chil. Hist. Nat. 77:411–437.
Eddington GE, Robbins WW. 1920. Irrigation water as a factor in the dissemination of weed seeds. Colo
Exp. Sta. Bull. 253:25.
Fairbrothers DE, Gray, JR (1972) Microstegium vinineum (Trin.) A. Camus (Gramineae) in the United
States. Bull. Torrey Bot. Club 99:97–100.
Flint ML, Van Den Bosch R. 1981. Introduction to Integrated Pest Management. Springer Press, New York,
256 pp.
Harper JL. 1977. Population Biology of Plants. Academic Press, New York, 892 pp.
Holm LG, Plucknett DL, Pancho JV, Herberger JP. 1977. The World’s Worst Weeds: Distribution and
Biology. East-West Center/University Press of Hawaii, Hawaii, 609 pp.
King LJ. 1966. Weeds of the World, Biology and Control. Interscience Publishers, New York,
270 pp.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

20 WEED BIOLOGY AND CLIMATE CHANGE

Koch W. 1969. Influence of environmental factors on the seed phase of annual weeds, from the point of
view of weed control. Habilitations-schrift Landwehrshaft, 50:204–210.
Kropff MJ, Spitters CJT. 1991. A simple model of crop loss by weed competition from early observations
on relative leaf area of the weeds. Weed Res 31:97–105.
Labrada R. 2007. The need for improved weed management in rice. In: Proceedings of the 20th Ses-
sion of the International Rice Commission, Bangkok, Thailand, July 23–26, FAO, Rome, pp. 310–
324.
Miller JH. 2003. Nonnative Invasive Plants of Southern Forests: A Field Guide for Identification and Control.
USDA, Forest Service, General Technical Report SRS-62, 93 pp.
Mohan JE, Ziska LH, Schlesinger WH, Thomas RB, Sicher RC, George K, Clark JS. 2006. Biomass and
toxicity responses of poison ivy (Toxicodendron radicans) to elevated atmospheric CO2 . Proc. Natl.
Acad. Sci. 103:9086–9089.
Monaco TJ, Weller SC, Ashton FM. 2002. Weed Science: Principles and Practices. John Wiley & Sons,
New York, 671 pp.
Mooney HA, Hobbs RJ. 2000. Invasive Species in a Changing World. Island Press, Washington, DC, 457
pp.
Novak SJ, Mack RN. 2001. Tracing plant introduction and spread: Genetic evidence from Bromus tectorum
(Cheatgrass). Bioscience 51:114–122.
Odum S. 1965. Germination of ancient seeds: Floristical observations and experiments with archaeologically
dated soil samples. Dan. Bot. Ark. 24:2.
Parsons WT. 1992. Noxious Weeds of Australia. Inkata Press, Melbourne, Australia.
Patterson DT. 1995. Effects of environmental stress on weed/crop interactions. Weed Sci. 43:483–
490.
Probert RJ. 1992. The role of temperature in germination ecophysiology. In: Fenner M (ed.) Seed: The
Ecology of Regeneration in Plant Communities. C.A.B. International, Wallingford, UK, pp. 285–
325.
Proctor VW. 1968. Long-distance dispersal of seeds by retention in digestive tract of birds. Science
160:321–322.
Radosevich S, Holt J, Ghersa C. 1997. Weed Ecology: Implications for Weed Management. John Wiley &
Sons, New York, 589 pp.
Scott HD, Geddes RD. 1979. Plant water stress of soybean (Glycine max) and common cocklebur (Xanthium
pennsylvanicum): A comparison under field conditions. Weed Sci. 27:285–289.
Shafroth PB, Cleverly JR, Dudley TL, Taylor JP, Van Riper C, Weeks EP, Stuart JN. 2005. Control
of Tamarix in the Western United States: Implications for water salvage, wildlife use, and riparian
restoration. Environ. Manag. 35:231–246.
Sharkey TD, Loreto F. 1993. Water stress, temperature, and light effects on the capacity for isoprene
emission and photosynthesis of kudzu leaves. Oecology 95:328–333.
Skinner K, Smith L, Rice P. 2000. Using noxious weed lists to prioritize targets for developing weed
management strategies. Weed Sci. 48:640–644.
Sauer J, Struik G. 1964. A possible ecological relation between soil disturbance, light flash, and seed
germination. Ecology 45:884–886.
Stallings GP, Thill DC, Mallory-Smith CA, Lass LW. 1995. Plant movement and seed dispersal of Russian
thistle (Salsola iberica). Weed Sci. 43:63–69.
Teasdale JR, Mohler CL. 1993. Light transmittance, soil temperature, and soil moisture under residue of
hairy vetch and rye. Agron. J. 85:673–680.
Vengris J, Colby WG, Drake M. 1955. Plant nutrient competition between weeds and corn. Agron. J.
47:213–216.
Westbrooks R. 1998. Invasive Plants, Changing the Landscape of America: Fact Book. Federal Interagency
Committee for the Management of Noxious and Exotic Weeds (FICMNEW), Washington, DC, 109
pp.
P1: SFK/UKS P2: SFK
c01 BLBS066-Ziska October 13, 2010 15:23 Trim: 244mm X 172mm Printer Name: Yet to Come

A BRIEF HISTORY OF WEEDS AND THEIR IMPACT 21

Wilson RG. 1980. Dissemination of weed seeds by surface irrigation water in western Nebraska. Weed Sci.
28:87–92.
Ye-Ebiyo Y, Pollack RJ, Spielman A. 2000. Enhanced development in nature of larval Anopheles arabiensis
mosquitoes feeding on maize pollen. Am. J. Trop. Med. Hyg. 63:90–93.
Zhang ZP. 2001. Weed management in rice in China. In: Summary Presented at FAO Workshop on
Echinochloa spp. Control, Beijing, China, May 27, 2001.
Zimdahl RL. 1993. Fundamentals of Weed Science. Academic Press, New York, 454 pp.
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

2 Carbon Dioxide and Global Warming: The “Green”

in the Greenhouse Effect

“Yes, it is real, sure is.”

George W. Bush in an interview with the Politico answering the question, “For the
record, is global warming real?”, May 13, 2008

Introduction

It may be hard to believe, but the air you breathe in today, even in the most remote parts of the
world, differs from the air that you breathed on the day you were born. Although you do not
notice the difference, the plants around you, whether they are crops or weeds or native plants
growing in the wild, certainly do. This chapter focuses on the atmosphere—primarily the
change in the concentration of one gas, carbon dioxide—and the ways that the environment
is responding. Put briefly, the climate is changing and plants are not growing quite like they
used to. Why?

The Greenhouse Effect Made Simple!

For all practical purposes, Earth’s climate depends on energy that arrives from the sun, and
what happens to that energy once it reaches our atmosphere. If the sun’s energy accumulated
continuously on Earth, the planet would get warmer and warmer. In fact, our planet has
reached, for now, a relatively stable temperature balance, where the energy coming in from
the sun is balanced by energy leaving. Over tens and hundreds of thousands of years, this
balance changes a bit, in what have been fairly regular cycles. For our purposes, though,
these longer-term fluctuations are less relevant than what keeps the planet at its current
temperature.
The sun is not the only object in space that sends energy our way. In fact, Earth receives
energy from anything in space that is warmer than absolute zero (although on balance it loses
energy to things cooler than itself). The warmer an object is, the more energy it loses to its
surroundings via electromagnetic radiation, as per the Stefan–Boltzmann law. Because the
sun is so hot, it gives off a tremendous amount of energy, and we benefit.

Weed Biology and Climate Change Lewis H. Ziska and Jeffrey S. Dukes
© 2011 Blackwell Publishing Ltd. ISBN: 978-0-813-81417-9

23
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

24 WEED BIOLOGY AND CLIMATE CHANGE

However, Earth also loses energy to space and that energy departs in its own spectrum of
wavelengths. Because Earth is so much cooler (relative to the sun), less energy is lost, and
the energy that departs has, on average, much longer (and thus less powerful) wavelengths.
The departing energy falls into the infrared portion of the electromagnetic spectrum and is
thus susceptible to absorption by greenhouse gases. The atmosphere’s ability to trap infrared
wavelengths—basically heat—depends on its composition. More of a given greenhouse gas
will mean more energy absorption at the specific wavelengths absorbed by that gas. As
different types of greenhouse gases enter the atmosphere, they increase energy absorption at
a wider range of wavelengths. More energy absorption by the atmosphere means a warmer
atmosphere, because less energy is lost to space. The system is sometimes described as a
greenhouse, in that light can penetrate the greenhouse, but that the glass (i.e., atmosphere)
prevents the long-wave infrared radiation (heat) from escaping. This is not, in itself, detri-
mental to biology; indeed, without this “greenhouse effect” the average temperature of the
earth would be about −18◦ C!

What Is a Greenhouse Gas?

It turns out that many gases can absorb infrared wavelengths of light. The most common of
these is water vapor. Others include carbon dioxide, methane, ozone, and many refrigerants.
Some of these gases play a much larger role in changing the climate than others. The
importance of a gas for climate change can depend on many factors such as how fast humans
are releasing it, how much is already in the atmosphere, how long the gas stays in the
atmosphere, and whether other gases already absorb the same wavelengths of energy.
Water vapor is the most abundant greenhouse gas, and humans have little influence on its
atmospheric concentration. Water vapor varies dramatically from place to place; the warm
air of the equatorial tropics can hold 10–20 times more water vapor than air at the poles.
When it comes to changing the climate, CO2 is the most important greenhouse gas,
primarily because we have changed its abundance in the atmosphere so dramatically—we
have stuffed an additional 200+ billion metric tons of carbon, in the form of CO2 , into an
atmosphere that used to hold less than 600 billion.
Several other greenhouse gases play important roles in climate change, including methane,
nitrous oxide, and halocarbons (such as chlorofluorocarbons, or CFCs). Their relative im-
portances are displayed in Table 2.1.

Where Do These Greenhouse Gases Come From?

Most of the major greenhouse gases were present in the atmosphere millions of years before
humans appeared on Earth. However, people have dramatically changed the cycling of these
gases. For example, every year, vast quantities of CO2 are taken up by plants on land and
by phytoplankton in the ocean. The carbon has been “fixed” by these organisms—integrated
into carbohydrates and other compounds—for some period of time. As these organisms
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 25

Table 2.1 “Radiative forcing” values (RFs) for the

most important greenhouse gases.
Common name Radiative forcing (W m−2 )

Carbon dioxide 1.56

Methane 0.59
Halocarbons 0.34
Nitrous oxide 0.15

These values describe how strongly the total emissions

of these gases since 1750 were changing the climate in
2005. Data from Forster et al. (2007).

die and decompose, or as they use carbohydrates to power their metabolic processes, they
release some of that carbon back to the atmosphere, again as CO2 . At the same time, CO2 is
continually dissolving into and back out of the ocean through physical processes. All these
cycles are remarkably well balanced, leading to relatively stable CO2 concentrations in the
atmosphere over at least the last 800,000 years (Lüthi et al., 2008; Siegenthaler et al., 2005).
However, during the last approximately 500 million years, some of the dying plants and
phytoplankton have not fully decomposed, instead getting buried into a geological limbo of
sorts, where time, pressure, and heat have incorporated some of their carbon into coal, oil,
or natural gas (Dukes, 2003). This stored carbon largely powered the Industrial Revolution
and the subsequent transformation of society, and its combustion has vented more than 300
billion tons of carbon into the air, principally as carbon dioxide (Canadell et al., 2007).
Agriculture and other activities have released another ∼158 billion tons of carbon formerly
stored in soils and plants (Canadell et al., 2007). Much of this was released from forests and
forest soils as areas were converted to agricultural or grazing lands. These conversions, now
primarily in tropical countries, still account for about 15–20% of all CO2 releases worldwide
(Canadell et al., 2007). Interestingly, the CO2 released to the atmosphere by human actions
has not all stayed there. We will discuss where this CO2 has gone later in this chapter.
As with CO2 , the concentration of atmospheric methane (CH4 ) has rapidly increased over
the last 150 years, more than doubling during this time. Interestingly, the rate of growth
has slowed dramatically since the early 1990s and the concentration may be approaching
equilibrium with current emission rates (Forster et al., 2007). Methane is produced and
consumed by widespread and common classes of microbes. The greatest rates of microbial
production take place in wet (especially anaerobic) environments. Conversion of large areas
to wet environments such as paddies for rice agriculture has increased methane production on
the land surface. The growth of the livestock industry has also increased methane production
because methane is generated in the digestive systems of ruminants. Methane is also released
during the mining and distribution of fossil fuels, from landfills, and when biomass is burned.
Microbes in the soils and the ocean naturally emit nitrous oxide (N2 O) as they conduct
nitrification and denitrification, but human activities have increased production of this gas
by approximately 18% over preindustrial levels (Forster et al., 2007). Fertilized agricultural
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

26 WEED BIOLOGY AND CLIMATE CHANGE

fields, especially when wet and when plants are taking up little or no nitrogen (such as when
plants are still small or when crops have not yet been planted), provide ideal conditions for
microbes to rapidly convert dissolved nitrogen into gaseous forms, including N2 O. Rapid
denitrification also occurs when dissolved nitrogen runs off from these fields, and in manure
management systems for livestock (Monteny et al., 2005; Stehfest and Bouwman, 2006).
Other processes that create substantial amounts of N2 O include fossil fuel combustion,
biomass burning, and some industrial processes such as nylon manufacturing (Denman
et al., 2007).
Human actions are simultaneously destroying ozone in the stratosphere (the region from
approximately 10 to 48 km in altitude) and stimulating its production in the troposphere
(the atmosphere beneath the stratosphere). Different processes are operating at the different
altitudes and the changes in concentration of this greenhouse gas have adverse health effects
for plants (and humans) at each level. In the stratosphere, ozone molecules absorb ultraviolet
(UV) radiation. As this ozone is destroyed through interactions with CFCs and other chemi-
cals, more of the harmful radiation reaches the earth’s surface, damaging nucleic acids and the
photosynthetic apparatus. Consequently, UV damage may lead to decreased photosynthesis
and plant growth (Caldwell et al., 2007). In humans, UV damage may lead to skin cancer. In
the troposphere, releases of nitrogen oxides, volatile organic compounds, and other gases,
often as products of fossil fuel combustion, increase the rate of ozone production. Ozone
at Earth’s surface enters leaves and stems and reacts with chemical constituents of plants,
altering their biochemistry in ways that eventually reduce the growth of plant (Ashmore,
2005). In humans, ozone intensifies breathing ailments (asthma, emphysema, etc.). At all
altitudes, ozone acts as a greenhouse gas.
Many synthetic halocarbons, including the CFCs that destroy ozone, are themselves pow-
erful greenhouse gases, although they exist only in small concentrations in the atmosphere.
Unlike the other greenhouse gases, most of the halocarbons are entirely manmade, for use as
refrigerants, propellants, and other purposes. Production of many halocarbons has decreased
or been halted following the passage of the Montreal Protocol, which was designed to phase
out the manufacture of chemicals that led to depletion of stratospheric ozone.

Measuring Greenhouse Gases

The story of the realization that humans were changing the climate is well described by Weart
(2003) and by Christianson (1999). In the 1950s, there was still some question about whether
the carbon dioxide coming out of smokestacks and tailpipes would stay in the atmosphere.
By comparing the isotopes, or kinds, of carbon in wood from old trees with those in new
wood, the chemist Hans Seuss found that vegetation had taken up some of the newly released
“fossil” carbon dioxide. It seemed quite possible—likely, even—that the vast majority of
the carbon dioxide released by humans had dissolved in the ocean, leaving the atmosphere
relatively unchanged. But by examining, among other things, the various isotopes of carbon
present in the atmosphere, Seuss and his colleague Roger Revelle found evidence that the
ocean was not rapidly taking up CO2 , and would only undo any changes in the atmosphere
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 27

Figure 2.1 Atmospheric carbon dioxide concentration as measured at the monitoring station on Mauna Loa, Hawaii.
This graph is also known as the “Keeling Curve.” Collection of data continues today at this site. The annual fluctuations
occur as a result of the seasons on Earth, and the greater land mass in the northern hemisphere. As plants take up carbon
dioxide in the northern hemisphere during the spring and summer, atmospheric CO2 concentrations decline. As plants in
the northern hemisphere go dormant in fall and winter, CO2 concentrations rise quickly. The accelerating long-term rise
in the graph is attributable to fossil fuel burning and land use change. Throughout the record of monitoring on Mauna
Loa, researchers have taken great care to factor out any gas samples contaminated with carbon dioxide from volcanic
gases. Based on data from Keeling et al. (2009).

over many generations. Shortly after, Revelle hired a young atmospheric chemist, Charles
Keeling, who might be able to show whether concentrations of the gas were increasing.
Keeling established the world’s first long-term, highly sensitive CO2 monitoring station
in Hawaii, on the volcanic slopes of Mauna Loa. Data from this station now make up one
of the most famous graphs in environmental science, known as the “Keeling Curve,” which
demonstrates how rapidly human activity has changed the atmosphere (Figure 2.1).
Initially, detecting an increase in atmospheric carbon dioxide concentrations was not a
simple task; in the 1950s, no company made scientific instruments sensitive enough to detect
CO2 in the parts per million (ppm) range. Charles Keeling had taken it upon himself to better
understand why Earth’s atmosphere had so little of a gas that had such an important role in
life. At CalTech, in 1955, he built the most sensitive instrument in existence to measure CO2 ,
and then carried out an obsessive spree of measurements. He started by carrying vacuum-
containing glass spheres up to the roof every 4 hours, holding his breath, and then opening the
flasks briefly, allowing them to fill with a sample of the region’s air. This continued for days
on end, during which he would leave his pregnant wife in the middle of the night to maintain
his schedule of measurements. Soon afterward, he brought his patient wife and infant son
into the mountains to camp in remote locations, as he again regularly filled flasks with air
(here uncontaminated by the urban Pasadena environment). This obsession served him well.
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

28 WEED BIOLOGY AND CLIMATE CHANGE

After a couple of years, Revelle offered him a job at the Scripps Institution of Oceanography
(where Seuss was also working). In his new position, Keeling was encouraged to follow his
passion and to establish the first CO2 monitoring stations.

It is interesting to note that Keeling’s most valuable measurements almost did not
get taken. Initial funding for Keeling’s CO2 station in Hawaii came from a year-long
international campaign to measure geophysical processes, and Keeling’s funding over
the following 5 years was generously redirected from grants awarded to Revelle and
Seuss at Scripps. Finally, though, the well ran dry. Government funding agencies
were reluctant to fund a monitoring project to address a question that, at least until
recently, was thought to be essentially irrelevant; after all, how could man’s activities
possibly change something so vast as Earth’s atmosphere? Fortunately, Keeling finally
convinced the U.S. National Science Foundation to fund his monitoring station in
1964, and the program resumed—leaving only a short gap in the record, and paving
the way for a crucially important data set. If not for the establishment and maintenance
of this monitoring project, the dramatic and ongoing impact of human activity on the
atmosphere might not have been recognized for several more years.

Follow the Carbon (Where Will It Take Us?)

Today, many human actions increase atmospheric carbon dioxide concentrations. Most of the
CO2 released in industrial nations comes from combustion of coal, oil, and natural gas, which
generates electricity and heat for buildings and which powers most transportation. A map of
carbon dioxide releases from the United States shows how, at the local scale, much of the
carbon dioxide is released from cities and roadways, as well as from more remote fossil–fuel
burning power plants and industrial plants (Figure 2.2; Gurney et al., 2009). However, another
source of carbon comes from the tropics, where conversion of forests to agricultural land
results in massive carbon dioxide emissions as trees are burned or decompose. These “land use
changes” account for approximately one-fifth of human-generated CO2 releases (Canadell
et al., 2007). All this carbon dioxide mixes relatively rapidly in the atmosphere.
The rate of carbon dioxide release has accelerated, almost continuously, since the begin-
ning of the Industrial Revolution. The concentration had hovered at or below 280 ppm for at
least the last 800,000 years, but as you read this, it has increased to about 390 ppm. This is
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 29

(log Kilotonne C/Year)

–5.7 to –0.4
–0.3 to 0.1
0.2 to 0.5
0.6 to 0.9
1.0 to 3.9

Figure 2.2 Spatial distribution of carbon dioxide release in the United States. Reprinted from Gurney et al. (2009).
Available in color online at http://www.purdue.edu/eas/carbon/vulcan/plots.php.

probably the highest atmospheric CO2 concentration of the last 20 million years (Canadell
et al., 2007).
What drives this acceleration? Increasing human population, economic activity, and de-
pendence on fossil fuel energy have acted together to accelerate CO2 releases. In addition,
population growth and economic pressures in developing countries have led to conversion
of tropical forests to agricultural land. The “cooking” of limestone for cement manufac-
ture also contributes a measurable fraction of human-generated CO2 , both because carbon
dioxide is released from the limestone as it is heated and because the heating itself is an
energy-intensive process. As population and world economies grow, demands for energy,
forest conversion, and cement continue to rise (Figure 2.3; Boden et al., 2009; Houghton,
2008). How fast society releases CO2 in the future will depend on several things, including
the rate of human population growth and the rate at which economic growth fuels growth in
energy consumption, but also the rate at which societies shift away from using fossil fuels
and the rate at which energy efficiency increases. Each of these rates is related to many
other factors in society, such as the rate of technological development, the dependence of
nations on energy produced in other countries, and political realities and implications sur-
rounding this dependence, the market prices of fossil fuels, and so on. Thus, CO2 releases are
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

30 WEED BIOLOGY AND CLIMATE CHANGE

Figure 2.3 Rate of human-generated carbon emissions from fossil fuel burning, cement production, and changing land
use since 1850. Based on data from Boden et al. (2009) and Houghton (2008).

intricately tied to many societal activities and decisions and this makes the regulation of CO2
emissions a major political and logistical challenge.
To our great fortune, not all the CO2 that we send into the atmosphere stays there. More
than half of this greenhouse gas is taken up on land or at sea, thanks to biological and
chemical processes. Carbon dioxide can dissolve in water and the oceans absorb one-quarter
of the human-released CO2 , contributing among other things to their increasing acidity. Many
marine organisms that form shells or other structures from carbonates, such as shellfish and
corals, are threatened by these chemical changes (Doney et al., 2009). In contrast, carbon
dioxide is also an essential ingredient for plant photosynthesis and growth (along with water,
nutrients, and sunlight), and the increase in atmospheric CO2 concentration, along with
several other factors, has led to increased growth of plants on land. The additional carbon
storage in these plants and in the soils beneath them accounts for another ∼30% of the human-
released CO2 (Canadell et al., 2007). So, together, the oceans and ecosystems on land slow
the growth in atmospheric CO2 concentration to less than half the rate of CO2 release.
The future composition of the atmosphere, then, depends not only on the rate of CO2
release from human society, but also on the rate of CO2 uptake by Earth’s surface. Whether
this rate of uptake will continue in the future is not certain—a variety of biological processes
leading to uptake could conceivably saturate or slow in the coming decades. In recent years,
uptake by Earth’s surface has not kept pace with emissions, so the fraction of CO2 that stays
in the atmosphere has risen slightly, from about 40% in 1960 to approximately 45% today.
This is primarily because the fraction of emitted CO2 taken up by the oceans has declined
(Canadell et al., 2007).
Given the complexity of the global carbon cycle, can we make any predictions about
carbon dioxide concentrations in a future atmosphere? The most authoritative group when
it comes to this general topic is the Intergovernmental Panel on Climate Change (IPCC).
The IPCC is a body of scientists from around the world who periodically assess the latest
knowledge about climate change. The consensus reports produced by this group are not
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 31

10
Actual emissions: CDIAC
Actual emissions: EIA

CO2 emissions (GtC y–1)

9 450 ppm stabilization
650 ppm stabilization
A1FI
8 A1B
A1T
A2
7 B1
B2

5
1990 1995 2000 2005 2010

Figure 2.4 Observed CO2 emissions according to two sources (black lines) compared with emissions scenarios
used by the IPCC (solid gray lines) and emissions trajectories to achieve specified CO2 concentration targets (dashed
lines). From Raupach et al. (2007). Updated version available online in color at http://www.globalcarbonproject.org/
carbonbudget/index.htm.

only vetted by many expert scientists at universities and research institutes, but also by
governments and their scientists. These detailed, careful reports won the IPCC a share of the
Nobel Peace Prize in 2007. In preparing the reports that examine how climate may change,
the IPCC relies on a set of future scenarios that describe how society may change over the
next 100 years, and how emissions of CO2 and other greenhouse gases would change in
each of those possible futures. Most of these scenarios are based on an assumption that, to
one degree or another, technological progress will help society to wean itself off fossil fuels
relatively quickly over the course of this century. The wide range of scenarios envisions that
atmospheric CO2 levels may rise to 463–623 ppm by 2050 and 478–1,099 ppm by 2100
(Carter et al., 2001). The highest emission scenario used by the IPCC is known as the A1FI
scenario, with “FI” abbreviating “fossil intensive.” Until the recent financial crisis, global
human activity was producing as much or more carbon dioxide than was envisioned under
this scenario, indicating that the IPCC’s most recent worst-case scenarios for climate change
could be somewhat optimistic (Figure 2.4; Raupach et al., 2007; Le Quéré et al., 2009).
Of course, carbon dioxide is not the only greenhouse gas affected by human actions. As
mentioned previously, several other greenhouse gases (methane and nitrous oxide, along with
ozone and CFCs) also have grown more concentrated in the atmosphere because of human
actions (Figure 2.5). Water vapor, too, is becoming more concentrated in the atmosphere
because of humans, but in this case the increase is a consequence of the addition of the
other greenhouse gases. As these other gases warm the atmosphere, water evaporates more
readily. Once in the atmosphere the additional water vapor acts as a potent greenhouse gas,
which may cause more warming than even the greenhouse gases initially released by humans.
More warming, in turn, leads to more water vapor. This phenomenon is known as a positive
feedback. The buildup of greenhouse gases is causing the atmosphere to trap heat more
effectively, warming the surface of the planet. But how much more will it warm, where will
it warm the most, and how do we know?
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

32 WEED BIOLOGY AND CLIMATE CHANGE

1,750 Methane 0.5

1,500 0.4

CH4 (ppb)
0.3
1,250
0.2
1,000
0.1
750 0.0

0.15
310 Nitrous oxide
0.10
N2O (ppb)

290
0.05

270 0.0

250
1000 1200 1400 1600 1800 2000
Year

Figure 2.5 Graph of increasing CH4 and N2 O concentrations. Atmospheric concentration is shown on the left axis,
and radiative forcing, or the relative influence of each gas on Earth’s energy balance, is shown on the right axis
(in W m–2 ). Figure from IPCC (2001).

Because the basic properties of greenhouse gases are known, we can estimate how well a
given atmosphere would insulate Earth’s surface from space. Clearly, increasing the concen-
tration of greenhouse gases should cause the surface to warm. How much it will warm is a
much more complicated question because of the many feedback loops in Earth’s climate such
as the positive feedback from water vapor. Another positive feedback loop may be related to
albedo or the amount of energy reflected back into space. For example, warming will mean
that less of the planet is covered by ice and snow. In turn, less of the sun’s energy will be
reflected back to space, and instead water, plants, and soil will capture that energy, warming
the planet further. A slightly warmer planet will mean that trees can grow at higher elevations
and latitudes, making portions of Earth’s surface darker still. Some other potentially impor-
tant feedbacks (positive and negative) such as the consequences of increased cloudiness are
less well understood.
Energy absorbed by Earth’s surface gets redistributed on a vast scale by ocean currents.
Winds and storm systems also rearrange large amounts of energy. Our knowledge about the
workings of the climate system, the properties of greenhouse gases, and the workings of
climate feedbacks are all necessary to predict how our actions will affect Earth’s climate.

Looking into the Crystal Ball: Climate Models and Projections

The best attempts by climate scientists to understand how differential changes in surface
temperatures might alter climate are based on complex mathematical representations of the
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 33

physical world: models that simulate the workings of the climate system on high-performance
computers. When the IPCC issues reports containing “climate projections,” these reports are
their best attempts to summarize what has been seen in these crystal balls. (Climate modelers
do not use the term “prediction” because they do not know what the actual rate of emissions
will be; each projection is essentially a prediction for the future if a certain set of conditions
comes to pass.)
Researchers in many different countries have created dozens of climate models over the
years, each with its own particular design and strategy for representing the climate system
and its various feedbacks. As more of the Earth system’s processes have been incorporated
into these models, the models have become more accurate at simulating past climates. The
IPCC summarizes results from the broad spectrum of climate models, across a variety of
future emission scenarios. While these models typically differ in their specific projections, the
general forecasts for warming and, in many areas of the world, precipitation, are remarkably
consistent (Solomon et al., 2007).
Under all the emission scenarios, the models foresee warming over the rest of this cen-
tury. The rate of warming depends on the emission scenario, although models project
relatively similar amounts of warming in the next few decades across diverse scenarios.
However, later in the century, different emission rates make major differences in projected
climates.
There are some general consistencies among models. First, warming will be greatest over
the land, as opposed to the ocean. This means that the estimates of “average” warming for
the globe reported in the media are typically lower than people and plants would actually
experience. Second, warming will be greatest at the poles, most dramatically at high northern
latitudes. Such projections are consistent with many of the recent data regarding polar
warming, increasing desertification, warmer winters, etc. Although future emission rates
will determine the intensity of warming, the spatial pattern of warming is consistent across
different scenarios (Figure 2.6; Meehl et al., 2007).
Climate models project shifts in precipitation patterns as well as temperature. In general,
precipitation is projected to increase at high latitudes, particularly in the winter. Many regions
of the world that are currently relatively dry are projected to become drier. These regions,
including the Mediterranean, southern Africa, and western Mexico, could be subject to
increased desertification (Figure 2.7). Globally, precipitation is projected to come in fewer
events per year; basically, storms will be more intense and arrive less often. In most regions,
this is bad news from the perspective of a plant; effectively, less of the precipitation that falls
will be available because more will run off the land during large storms. As the time between
storms increases, short droughts may become more frequent.
The climate models considered by the IPCC have a fairly low spatial resolution, largely
because the models demand so much computing power to run. Unfortunately, this coarse
scale prevents the models from accurately capturing effects of many local features such as
mountain ranges. Two techniques are used most often to create climate projections with local
detail. Regional climate models have been created to simulate climate in particular regions,
typically at the subcontinental scale. Statistical “downscaling,” working at similar scales,
interprets the output of the global climate models in the context of the typical local climate
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

34 WEED BIOLOGY AND CLIMATE CHANGE

Figure 2.6 Projected warming of Earth’s surface (relative to temperatures in 1980–1999) under scenarios of lower,
intermediate, and higher greenhouse gas releases. Until recently, the actual emission rate exceeded that used in the highest
release scenario. Note the consistent spatial variation in projected warming. Figure from Meehl et al. (2007). Available
in color online at http://www.ipcc.ch/publications and data/ar4/wg1/en/tssts-5-2.html

Projected change Projected change

Multi-model (in 2090–2099 for A1B) DJF multi-model (in 2090–2099 for A1B) JJA
20
©IPCC 2007: WG1-AR4

10
5
–5
–10
–20

Figure 2.7 Projected seasonal precipitation changes (relative to temperatures in 1980–1999) under a scenario of
intermediate greenhouse gas releases. The figures show the mean responses across several climate models. Areas with
black dots indicate that more than 90% of the models used agree on the nature of the change. The left panel shows changes
in December through February and the right panel shows changes in June through August. Figure from Solomon et al.
(2007). Available in color online at http://www.ipcc.ch/publications and data/ar4/wg1/en/tssts-5-3.html
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 35

relationships seen within a region. The products of these regional approaches are particularly
useful when contemplating how particular ecosystems may change in the future and they
provide opportunities for people in the region to consider how climate change will affect
them.

A Plant’s “Eye” View of the Future

How will plants respond to these changing conditions? This is not a trivial question, since
plants are the basis of the global food chain and any change in their biology will impact all
living systems. For obvious reasons, therefore, questions regarding plant function have been
the subject of thousands of scientific papers and talks. But as many researchers have deter-
mined, assessing the impact of climate change on plant biology is difficult, particularly on a
large scale. First of all, people are changing many aspects of the environment simultaneously.
The gas that plants use as a building block, carbon dioxide, has become nearly 40% more
abundant in just over a century. Imagine how we would respond if oxygen were 40% more
abundant! (Well, we might have massive problems with wildfires, but certainly we would
all be better athletes!) We are moving plants and their diseases, insect pests, and pollinators
from continent to continent around the world, so that plant species are now interacting with
a different community than that in which they most recently evolved. We are appropriating
habitat and altering it to serve our needs, causing the effective sizes of many plant populations
to shrink (with the notable exceptions of crops and many weed species), and reducing the
amount of available habitat in which seeds might germinate and survive. On the other hand,
we are regularly disturbing vast areas of land along roadsides, train tracks, in cities, and in
other areas associated with human activity, killing or cutting the native vegetation in these
areas and creating conditions in which fast-growing weeds can thrive. We are fertilizing
the world, on purpose as well as accidentally, via the nitrogen-containing constituents of
exhaust. We are spraying vast tracts of land with a variety of herbicides to benefit crops and
other desired species. And, on top of all this, we are changing the climate. These changes
are not all happening at the same rate in the same locations, but no plants are escaping the
changing atmosphere. We may develop decent predictions about how plants will respond to
one of these variables, but how might a second variable affect that response?
It is also difficult to predict how plants as a group will respond to future conditions because
there are so many different species and vegetation types, some of which are particularly
difficult to study. Imagine, for example, trying to characterize how tropical rainforest will
respond to one of these perturbations, even in a single region. This would involve trying
to understand the aggregate response of hundreds of different tree species (not to mention
the other plants), many of which live for hundreds of years. How should one choose which
species to study, and which life stages? Even then, how would we know whether the plant’s
response might change a relationship with a herbivore or a pollinator, and whether that
changing relationship might have even more important consequences for the forest than the
plant response itself? And still, all this ignores the practical challenges of working in such
an environment.
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

36 WEED BIOLOGY AND CLIMATE CHANGE

Despite these and other challenges, researchers have made progress. Much is understood
about the basic physiological responses of plants to increased levels of atmospheric carbon
dioxide, to warming, and to changes in precipitation.
It was suspected early on that these physiological responses would depend on the type of
photosynthetic machinery used by a plant. A variety of different photosynthetic pathways
are found in the plant kingdom, which can be loosely categorized as C3 , C4 , and CAM
photosynthesis—terms derived from the initial products of carbon fixation in each path-
way. C4 species like sugarcane and corn are mostly found in hot, dry environments, where
their physiology gives them a few competitive advantages. For one thing, they use water
efficiently—for each milliliter of water they lose through their leaves, they can grow more
than plants using the C3 pathway. Second, at high temperatures, they typically carry out
photosynthesis faster than C3 plants. Our changing atmosphere is identifying (or creating)
a drawback of their photosynthetic system, though: It is less responsive to high CO2 con-
centrations than that of C3 species. From these basic properties, we might expect warming
to favor C4 species, and increasing CO2 to favor C3 s. So, how will these different types of
species fare, in a warmer, more CO2 -rich world? Which species will be more competitive in
the future? We will discuss this more in the next chapter.
More will determine a species’ future success than its photosynthetic pathway, of course.
Many plants respond to environmental changes by altering their allocation to different tissue
types and changing their internal chemistry. For instance, plants grown in elevated CO2
typically have lower concentrations of nitrogen-containing proteins in their leaves. This
has consequences for the species that eat them, as these animals (or people) will get less
protein per mouthful in the future. Other chemical changes, such as altered levels of plant
toxins, could also change plant–animal interactions. Some of these chemical changes may
affect the way that weed species respond to management techniques such as the application
of herbicides. Changes in allocation to roots and shoots could also affect management
techniques. We will explore the possible consequences in the next chapter.

Summary

Human actions have injected vast quantities of greenhouse gases into the atmosphere over
the past approximately 150 years. These include CO2 , released primarily from combustion
of fossil fuels, methane, and nitrous oxide, released by agriculture and industry, and other
gases. Because they can absorb infrared radiation emitted from Earth’s surface, these gases
cause Earth’s atmosphere to warm up more than it previously could have. This, in turn, keeps
Earth’s surface warmer. Climate models suggest significant warming will occur by 2100,
particularly over land. This will be accompanied by changes in precipitation patterns. Plants
will respond directly to the increase in CO2 concentration, as well as to climate change and
other ongoing environmental changes. In the next two chapters, we will begin to address
in more detail the reported and potential changes in plant function resulting from climate
change, focusing in particular on weed biology.
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

CARBON DIOXIDE AND GLOBAL WARMING 37

References

Ashmore MR. 2005. Assessing the future global impacts of ozone on vegetation. Plant Cell Environ.
28:949–964.
Boden TA, Marland G, Andres RJ. 2009. Global, Regional, and National Fossil–Fuel CO2 Emissions.
Carbon Dioxide Information Analysis Center, Oak Ridge National Laboratory, U.S. Department of
Energy, Oak Ridge, TN.
Caldwell MM, Bornman JF, Ballaré CL, Flint SD, Kulandaivelu G. 2007. Terrestrial ecosystems, increased
solar ultraviolet radiation, and interactions with other climate change factors. Photochem. Photobiol.
Sci. 6:252–266.
Canadell JG, Le Quéré C, Raupach MR, Field CB, Buttenhuis ET, Ciais P, Conway TJ, Gillett NP, Houghton
RA, Marland G. 2007. Contributions to accelerating atmospheric CO2 growth from economic activity,
carbon intensity, and efficiency of natural sinks. Proc. Natl. Acad. Sci. U. S. A. 104:18866–18870.
Carter TR, La Rovere EL, Jones RN, Leemans R, Mearns LO, Nakicenovic N, Pittock AB, Semenov SM,
Skea J. 2001. Developing and applying scenarios. In: McCarthy JJ, Canziani OF, Leary NA, Dokken DJ,
White KS (eds) Climate Change 2001: Impacts, Adaptation, and Vulnerability. Cambridge University
Press, Cambridge, UK, pp. 145–190.
Christianson GE. 1999. Greenhouse: The 200-Year Story of Global Warming. Walker and Company, New
York.
Denman KL, Brasseur G, Chidthaisong A, Ciais P, Cox PM, Dickinson RE, Hauglustaine D, Heinze C,
Holland E, Jacob D, Lohmann U, Ramachandran S, Dias PLDS, Wofsy SC, Zhang X. 2007. Couplings
between changes in the climate system and biogeochemistry. In: Solomon S, Qin D, Manning M, Chen
Z, Marquis M, Averyt KB, Tignor M, Miller HL (eds) Climate Change 2007: The Physical Science
Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental
Panel on Climate Change. Cambridge University Press, Cambridge, UK, pp. 499–587.
Doney SC, Fabry VJ, Feely RA, Kleypas JA. 2009. Ocean acidification: the other CO2 problem. Ann. Rev.
Marine Sci. 1:169–192.
Dukes JS. 2003. Burning buried sunshine: human consumption of ancient solar energy. Clim. Change
61:31–44.
Forster P, Ramaswamy V, Artaxo P, Berntsen T, Betts R, Fahey DW, Haywood J, Lean J, Lowe DC, Myhre
G, Nganga J, Prinn R, Raga G, Schulz M, Dorland RV. 2007. Changes in atmospheric constituents and
in radiative forcing. In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt KB, Tignor M,
Miller HL (eds) Climate Change 2007: The Physical Science Basis. Contribution of Working Group
I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge
University Press, Cambridge, UK, pp. 129–234.
Gurney KR, Mendoza DL, Zhou Y, Fischer ML, Miller CC, Geethakumar S, de la Rue du Can S. 2009.
High resolution fossil fuel combustion CO2 emission fluxes for the United States. Environ. Sci. Tech.
43:5535–5541.
Houghton RA. 2008. Carbon flux to the atmosphere from land-use changes: 1850–2005. In: Trends: A
Compendium of Data on Global Change. Carbon Dioxide Information Analysis Center, Oak Ridge
National Laboratory, U.S. Department of Energy, Oak Ridge, TN.
Keeling RF, Piper SC, Bollenbacher AF, Walker JS. 2009. Atmospheric CO2 records from sites in the
SIO air sampling network. In: Trends: A Compendium of Data on Global Change. Carbon Dioxide
Information Analysis Center, Oak Ridge National Laboratory. U.S. Department of Energy, Oak Ridge,
TN.
Le Quéré C, Raupach MR, Canadell JG, Marland G, Bopp L, Ciais P, Conway TJ, Doney SC, Feely RA,
Foster P, Friedlingstein P, Gurney K, Houghton RA, House JI, Huntingford C, Levy PE, Lomas MR,
Majkut J, Metzl N, Ometto JP, Peters GP, Prentice IC, Randerson JT, Running SW, Sarmiento JL,
Schuster U, Sitch S, Takahashi T, Viovy N, van der Werf GR, Woodward FI. 2009. Trends in the
sources and sinks of carbon dioxide. Nature Geoscience 2:831–836.
P1: SFK/UKS P2: SFK
c02 BLBS066-Ziska October 7, 2010 9:5 Trim: 244mm X 172mm Printer Name: Yet to Come

38 WEED BIOLOGY AND CLIMATE CHANGE

Lüthi D, Floch ML, Bereiter B, Blunier T, Barnola J-M, Siegenthaler U, Raynaud D, Jouzel J, Fischer H,
Kawamura K, Stocker TF. 2008. High-resolution carbon dioxide concentration record 650,000–800,000
years before present. Nature 453:379–382.
Meehl GA, Stocker TF, Collins WD, Friedlingstein P, Gaye AT, Gregory JM, Kitoh A, Knutti R, Murphy
JM, Noda A, Raper SCB, Watterson IG, Weaver AJ, Zhao Z-C. 2007. Global climate projections. In:
Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt KB, Tignor M, Miller HL (eds) Climate
Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment
Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge,
UK, pp. 747–845.
Monteny G-J, Bannink A, Chadwick D. 2005. Greenhouse gas abatement strategies for animal husbandry.
Agric. Ecosyst. Environ. 112:163–170.
Raupach MR, Marland G, Ciais P, Quéré CL, Canadell JG, Klepper G, Field CB. 2007. Global and regional
drivers of accelerating CO2 emissions. Proc. Natl. Acad. Sci. U. S. A. 104:18866–18870.
Siegenthaler U, Stocker TF, Monnin E, Lüthi D, Schwander J, Stauffer B, Raynaud D, Barnola J-M,
Fischer H, Masson-Delmotte V, Jouzel J. 2005. Stable carbon cycle–climate relationship during the
late Pleistocene. Science 310:1313–1317.
Solomon S, Qin D, Manning M, Alley RB, Berntsen T, Bindoff NL, Chen Z, Chidthaisong A, Gregory JM,
Hegerl GC, Heimann M, Hewitson B, Hoskins BJ, Joos F, Jouzel J, Kattsov V, Lohmann U, Matsuno
T, Molina M, Nicholls N, Overpeck J, Raga G, Ramaswamy V, Ren J, Rusticucci M, Somerville
R, Stocker TF, Whetton P, Wood RA, Wratt D. 2007. Technical summary. In: Solomon S, Qin D,
Manning M, Chen Z, Marquis M, Averyt KB, Tignor M, Miller HL (eds) Climate Change 2007:
The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the
Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, UK, pp. 19–91.
Stehfest E, Bouwman L. 2006. N2 O and NO emission from agricultural fields and soils under natural
vegetation: summarizing available measurement data and modeling of global annual emissions. Nutr.
Cycling Agroecosyst. 74:207–228.
Weart S. 2003. The Discovery of Global Warming. Harvard University Press, Cambridge, MA.
P1: SFK/UKS P2: SFK
c03 BLBS066-Ziska October 13, 2010 17:50 Trim: 244mm X 172mm Printer Name: Yet to Come

3 An Evaluation of the Impact of Rising Carbon Dioxide

and Climatic Change on Weed Biology:
From the Cell to the Plant

“Though I do not believe that a plant will spring up where no seed has been, I have great
faith in a seed. Convince me that you have a seed there, and I am prepared to expect
wonders.”
Henry David Thoreau

Introduction

We have seen from the previous chapter that increases in carbon dioxide concentration
CO2 and the subsequent consequences for climate, particularly surface temperature and
hydrology, are associated with human activities, most notably the demand for energy (fossil
fuel consumption) and land (deforestation). Given these projected environmental changes,
what are the likely impacts as they relate to our understanding of the basic biology of weeds?
To answer this question, one needs to understand basic plant biology, how rising CO2 and
climate will alter plant biology on a number of different functional levels, and how those
levels interact with each other. To appreciate the complexity of plants, we must keep in mind
that biological processes at the cellular level (e.g., photosynthesis) happen quickly and at
very small scales, whereas those at the ecosystem level take longer and occur over larger
areas (e.g., plant-to-plant competition for resources, seed production) (Figure 3.1).
In this chapter, we examine these biological processes in detail and discuss how rising CO2
and the combination of rising CO2 and climate (precipitation and temperature) are likely
to impact weed biology. We consider different levels of functional organization from the
cellular level to the whole plant, using the construct as shown in Figure 3.1. The following
chapter will “scale-up” from the response of individuals to examine responses of plant–plant
interactions and biogeographical distribution.
This approach recognizes the certainty of continuing increases in global levels of atmo-
spheric CO2 , but also acknowledges that the extent to which the increase in CO2 and other
anthropogenic greenhouse gases alter surface temperature and water availability is likely to
vary temporally and geographically (Solomon et al., 2007).

Weed Biology and Climate Change Lewis H. Ziska and Jeffrey S. Dukes
© 2011 Blackwell Publishing Ltd. ISBN: 978-0-813-81417-9

39
Random documents with unrelated
content Scribd suggests to you:
 Ja – ja – es waren deutsche Feldgraue –, im heller werdenden
Morgenlicht sah er es nun deutlich.
Das war ein frohes Reiten! Durch den flachen Fluß ging's noch mit
einer letzten Anstrengung – dann hielt er unter den Seinen.
Man half ihm vom Pferd. Schwindlig, von Staub und Schweiß
unkenntlich, stand er zwischen den Soldaten, die ihn neugierig
umdrängten.
Atemlos fragte er nach dem Hauptmann Breuer, seine Meldung zu
machen. Man führte ihn schleunigst zu dem Offizier.
Die Ortschaft war genommen, der Russe vertrieben, so hörte
Helmut auf dem Wege. Zwar hatte der Feind noch einen Angriff vom
Walde her versucht, doch neu eintreffende Verstärkung aus der
Flanke hatten seinen Plan zuschanden gemacht.
Hauptmann Breuer hörte staunend auf Helmuts Bericht.
»Na, Junge – das war ein Meisterstück!« rief er und schüttelte
vergnügt Helmut die Hand. »Jetzt wollen wir den Kerls mal das
Erwachen versalzen. Gibt's denn nicht einen näheren Weg zu der
Scheune – ... Karte her ... Donnerschock, da sind ja unsere Leute
ganz in der Nähe – auf der anderen Seite vom Wald sollte das
Regiment auf weitere Befehle warten.«
»Kerls – Freiwillige vor! Wer rettet achtzig Kameraden?« Aus dem
Heidekraut sprang's empor – die Todmüden, vom harten Kampf des
Tages Erschöpften reckten sich auf. »Hier, Herr Hauptmann –!«
»Melde mich, Herr Hauptmann!« »Herr Hauptmann schicken Sie
mich!« Drei wurden ausgesucht – der Hauptmann zeigte auf der
Karte den Weg durch die brennende Ortschaft gerade hindurch, am
Flusse entlang war die deutsche Stellung in einer Stunde zu
erreichen! Statt des wohlverdienten Schlafes galt es einen Dauerlauf.
Die drei trabten tapfer davon!
Der Hauptmann rieb sich vergnügt die Hände! »So, mein Junge!
Oberst von Borwitz wird das übrige besorgen! Heut nachmittag
werden unsere Leute, denke ich, wieder bei uns eintreffen!«
»Herr Hauptmann, eine Bitte habe ich noch – können Sie nicht ein
paar Sanitäter nach dem Wald schicken? Dort liegt Herr von
Mansfeld verwundet, auch noch andere. Ich versprach's ihm ...«
»Gewiß – gewiß doch! Soll gleich geschehen! Hoffentlich ist der
gute Junge noch zu retten – seiner Mutter Einziger.
Na, und nun erst mal ran mit 'nem Schluck Rotwein und Brot und
Wurst! Weiter haben wir selber nichts!«
Der Hauptmann aber ließ Helmut die besten Teile seines
Frühstücks und schaute lachend zu, wie es dem schmeckte.
Am Nachmittag, wie Hauptmann Breuer es vorausgesagt, trafen
die achtzig Befreiten beim Bataillon ein. Es fehlte kein Einziger. Mit
einem kräftigen Hurra wurden sie von den Kameraden empfangen,
Helmut sprang seinem Vater an den Hals, jeder Schatten war
zwischen ihnen verschwunden.
Gegen Abend traten die Kompagnien am Ufer des Flüßchens zum
Appell an. Hell schien die warme Frühlingssonne dem Hauptmann
Breuer in das gute rotbraune Gesicht, als er die ihm anvertrauten
Mannschaften musterte. Ganz unten in der Reihe stand Helmut in
seinen zerlumpten Kleidern, eine feldgraue Mütze auf dem Kopf, der
Jüngste der Kompagnie.
Der Hauptmann hielt eine Ansprache.
»Leute!« sagte er, »ihr habt alle eure Pflicht ehrlich getan an dem
heißen Kampftag! Was uns befohlen war, das haben wir geleistet.
Aber achtzig von euch stünden heut nicht unter uns, wenn dieser
junge Zivilist hier nicht über die Köpfe der schnarchenden Feinde
hinweggestiegen wäre und durch einen kühnen Ritt ihnen die
Befreiung gebracht hätte! Helmut Kärn tritt vor!«
Militärisch stramm folgte der Junge dem Befehl.
 »Dir gebührt das Ehrenzeichen der Tapferkeit!«
Der Hauptmann löste das Eiserne Kreuz von seiner eigenen Brust
und heftete es Helmut an die zerrissene Jacke.
»Trage das Kreuz, den höchsten Schmuck des deutschen Soldaten
zur Erinnerung an deine mutige Tat! Bleibe durch dein ganzes Leben
seiner würdig! Ein Hurra unserem jungen Helden!«
»Hurra – Hurra – Hurra!« klang es dröhnend aus Hunderten von
rauhen Männerkehlen zum goldenen Abendhimmel empor.
Helmut stand stumm und zitternd vor Glück, während der
Hauptmann seinem Vater die Hand schüttelte. Dem rannen die
dicken Tränen in den zottigen Bart.
Dem Hauptmann Breuer war es immer wohler, wenn die
feierlichen Augenblicke vorüber waren, und er wieder gemütlich mit
seinen Leuten verkehren konnte. So nahm er denn, während die
Mannschaften abtraten, Helmut am Ohrläppchen und schüttelte ihn
lachend. »Verfluchter Bengel, jetzt geht's aber spornstreichs zurück
in die Schule! Auf die Hosen gesetzt und gebüffelt! Verstehst du
mich? Wehe dir, wenn's nächste Ostern keine Versetzung gibt!«
»Wird gemacht«, rief Helmut strahlend. »Nur eine Bitte noch, Herr
Hauptmann!«
»Na?«
»Daß Vater bald mal Urlaub kriegt!«
»Zugestanden! In acht Tagen ist er bei Muttern!«
 Wie Helmut nach Hause kommt

Die Fahrgäste der Elektrischen, welche die lange Schloßstraße in

Charlottenburg hinauffuhr, stießen sich an und machten sich
untereinander aufmerksam auf einen Jungen, der still in der Ecke
saß, und dessen tiefgebräuntes Gesicht immerfort glückselig in sich
hineinlachte. Zu einem neuen Anzug aus gutem kräftigen Stoff und
funkelnagelneuen Stiefeln trug er eine alte zerbeulte Feldmütze und
im Knopfloch seiner Jacke prangte schlicht und doch seltsam
eindrucksvoll das Eiserne Kreuz.
»Unverschämt, sich das anzuhängen, so'n Grünschnabel«,
schimpfte ein leberkrank aussehender Herr, »wer weiß, wo er das
gestohlen hat. Da sollte man doch den Schutzmann aufmerksam
machen – das ist grober Unfug!«
Richtig, als Helmut den Wagen verließ, um das letzte Stück zu
seiner Mutter Wohnung zu Fuß zu gehen, folgte ihm der grimmige
Junggeselle, trat an der Straßenecke auf einen Schutzmann zu und
flüsterte mit diesem.
Helmut bemerkte es nicht, er war ganz befangen von der
Erwartung des nahen Wiedersehens mit den Seinen. Die Offiziere
von seines Vaters Regiment hatten zusammengelegt, ihm die
Rückreise bezahlt und ihn in Königsberg mit neuen Kleidern
ausrüsten lassen. Nur von der alten Feldmütze, die ihm die
Kameraden des Vaters geschenkt, und die er in dem heiligsten
Augenblick seines Lebens getragen – von der konnte er sich nicht
trennen.
 Erschrocken blickte er sich um, als eine Hand sich auf seine
Schulter legte und eine bärbeißige Stimme fragte: »Sie, junger Herr,
wo haben Sie denn das Kreuz da her?«
»Das werde ich Ihnen gleich sagen, Herr Müller«, antwortet
Helmut stolz, denn der Schutzmann war ja sein alter Freund. Helmut
hatte ihn hundertmal um Auskunft gefragt, wenn er sich in dem
fremden Berlin nicht zurechtfand. Immer stand der Schutzmann mit
dem blanken Helm über dem friedlichen rosenroten Vollmondgesicht
an dieser Ecke auf treuer Wacht. Hinter seiner bärbeißigen Stimme
verbarg er ein freundliches, hilfsbereites Wesen und eine unendliche
Geduld.
»Ne nu seh einer – das ist ja der Brasilianer!« rief er jetzt lachend.
»Du Ausreißer – ganz Berlin haben wir nach dir durchsucht.... Wo
kommst du denn nun hergeschneit?«
Helmut zog aus seiner Brusttasche ein Papier und reichte es mit
heimlichem Schmunzeln der hohen Obrigkeit.
Inzwischen hatte sich ein Kreis von Neugierigen um die beiden
versammelt. Einige Frauen waren gleich hinter dem grilligen alten
Herrn aus dem Wagen gestiegen, um zu sehen, wie die Sache mit
dieser rätselhaften Person sich weiterentwickeln würde. Und da
gerade die Schule aus war, strömten die Jungen in hellen Haufen
herbei.
Der Schutzmann entfaltete weitläufig das Aktenstück, in dem der
Oberst des Regimentes Helmut Kärn bescheinigte, daß er das
Eiserne Kreuz sich mit Recht und Ehren erworben habe durch die
Rettung von achtzig deutschen Soldaten aus Feindeshand und durch
seinen dabei bewiesenen Mut. Der Schutzmann las aufmerksam von
Anfang bis zu Ende und betrachtete genau den Stempel des
Regiments.
»Na, denn wäre ja woll alles in Ordnung«, sagte er, Helmut das
Papier zurückgebend, »du bist ja ein Mordskerl! Aber trag' das Papier
man lieber immer bei dir! So 'ne verwunderliche Sache – die glaubt
sonst kein Mensch so 'nem Lausejungen! Meinen Glückwunsch!« Er
legte salutierend die Hand an den Helm. »Lesen, lesen – laut lesen!«
schrie es rings im Kreis von alten und jungen Stimmen. Helmut
überfiel plötzlich eine fürchterliche Verlegenheit. Es war doch gar
nicht leicht, sich zu Haus so richtig als Held zu benehmen. Er hielt
sein wertvolles Aktenstück fest in der Hand, machte sich mit Puffen
und Stößen heftig Bahn durch die andrängende Jungenschar und
rannte davon, so schnell ihn seine Beine trugen. Die Schuljungen mit
Hallo und Geschrei ihm nach. Wie ein junger Hirsch entsprang er
den Verfolgern, erreichte schweißtriefend das Haus, in dem die
Großeltern wohnten und war heilsfroh, als die schwere Eingangstür
hinter ihm ins Schloß fiel.
Dreimal riß er oben an der Wohnungsklingel, wie es seine Art war,
wenn er hungrig aus der Schule kam. Drinnen entstand ein Lärm,
ein Umstoßen von Stühlen, ein Rufen: »Das ist er – Helmut –
Helmut!«
Die Tür wurde aufgerissen, lachend und weinend küßte ihn seine
Mutter.
Hinter ihr zwischen der Gänseblume und der Großmutter erschien
noch ein anderes blühendes, braunäugiges Gesicht. Dort stand Frau
Anna Ledderhose, die der Mutter in der schweren Zeit treulich
geholfen hatte. Am Kaffeetisch bei all den lieben vertrauten
Menschen konnte Helmut nun nach Herzenslust erzählen und
berichten von seinem reichen, ernsten Erleben. Die Mutter hielt
seine Hand, die sie immer wieder streichelte und drückte. Nachher,
als sie beide allein waren, gab er ihr das Aktenstück und nahm das
Ehrenzeichen von seiner Jacke.
»Du, Mutti, verwahre es mir, bis ich erwachsen bin«, sagte er
»dann will ich's tragen. Jetzt so als ein Wundertier in Berlin
herumlaufen und in der Schule protzen – dazu ist mir das Eiserne
Kreuz zu heilig.«
Friedrich Andreas Perthes A.-G. Gotha
 Hinweise zur Transkription

Der Vermerk "Z. XI." auf der Titelseite ist ein Ausfuhrzeichen aus der
Weltkriegszeit.
Die ganzseitigen Illustrationen "Vater fährt ins Feld" (S. 25),
"Landarbeit im Kriege" (S. 43), "Ost-Preussen" (S. 55) und "Die
Flucht" (S. 91) wurden vor den Beginn des jeweiligen Kapitels
verschoben, um den Lesefluss nicht zu stören.
Das Originalbuch ist in Frakturschrift gedruckt.
Darstellung abweichender Schriftarten: g e s p e r r t , Antiqua.
Entsprechend der überwiegenden Darstellung im Originalbuch wurde
das Komma einheitlich hinter das schließende Anführungszeichen
gesetzt.
Der Text des Originalbuches wurde grundsätzlich beibehalten, mit
folgenden Ausnahmen,
Seite 8:
"«" eingefügt
(Wie alt sind Sie denn?«)
Seite 15:
"laß" geändert in "las"
(Der Lehrer las laut vor)
Seite 17:
"›" eingefügt
(›Morgen hast'n blauen Lappen dafor‹)
Seite 17:
"›" entfernt vor "Weg"
(Weg is er, und ick konnte nicht hinterher.)
Seite 42:
"den altem" geändert in "dem alten"
(mit dem alten Lodenmantel ihres Mannes angetan)
Seite 57:
"»" eingefügt
(»Wir haben uns nur im Walde versteckt)
Seite 64:
"«" eingefügt
(erklären Sie mir blos in aller Welt ...?«)
Seite 73:
"«" eingefügt
(eine Karte von ihm aus Sibirien.«)
Seite 77:
"." eingefügt
(laut aufs Knie und lachte.)
Seite 79:
"»" eingefügt
(»Siehst du, das alles hier soll mal)
Seite 79:
"göttlicheschönen" geändert in "göttlichschönen"
(Schmuck dieser köstlichen, göttlichschönen Geschichten)
Seite 79:
"»" entfernt vor "Nu"
(Nu – wir haben's ja dazu!)
Seite 81:
"überdem" geändert in "über dem"
(gleich einer schiefen Mütze über dem Gebäude)
Seite 82:
"," geändert in "."
(an der Schulter und schüttelte ihn.)
Seite 83:
"Herre nentfernten" geändert in "Herren entfernten"
(Auch die beiden jungen Herren entfernten sich)
Seite 85:
"Finde" geändert in "Finte"
(eine Finte von unserem Hauptmann)
Seite 85:
"«" entfernt hinter "Ende!"
(wenigstens hat die Warterei ein Ende!)
Seite 88:
"«" eingefügt
(bei den besoffenen Kerls durchschleichst ...«)
Seite 93:
"»" eingefügt
(»Melde mich, Herr Hauptmann!«)
Seite 93:
"»" eingefügt
(»Herr Hauptmann schicken Sie mich!«)
Seite 93:
"«" eingefügt
(wieder bei uns eintreffen!«)
Seite 94:
"«" eingefügt
(Ich versprach's ihm ...«)
 *** END OF THE PROJECT GUTENBERG EBOOK WAS HELMUT IN
DEUTSCHLAND ERLEBTE: EINE JUGENDGESCHICHTE ***

Updated editions will replace the previous one—the old editions will
be renamed.

Creating the works from print editions not protected by U.S.

copyright law means that no one owns a United States copyright in
these works, so the Foundation (and you!) can copy and distribute it
in the United States without permission and without paying
copyright royalties. Special rules, set forth in the General Terms of
Use part of this license, apply to copying and distributing Project
Gutenberg™ electronic works to protect the PROJECT GUTENBERG™
concept and trademark. Project Gutenberg is a registered trademark,
and may not be used if you charge for an eBook, except by following
the terms of the trademark license, including paying royalties for use
of the Project Gutenberg trademark. If you do not charge anything
for copies of this eBook, complying with the trademark license is
very easy. You may use this eBook for nearly any purpose such as
creation of derivative works, reports, performances and research.
Project Gutenberg eBooks may be modified and printed and given
away—you may do practically ANYTHING in the United States with
eBooks not protected by U.S. copyright law. Redistribution is subject
to the trademark license, especially commercial redistribution.

START: FULL LICENSE

THE FULL PROJECT GUTENBERG LICENSE
 PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg™ mission of promoting the free

distribution of electronic works, by using or distributing this work (or
any other work associated in any way with the phrase “Project
Gutenberg”), you agree to comply with all the terms of the Full
Project Gutenberg™ License available with this file or online at
www.gutenberg.org/license.

Section 1. General Terms of Use and

Redistributing Project Gutenberg™
electronic works
1.A. By reading or using any part of this Project Gutenberg™
electronic work, you indicate that you have read, understand, agree
to and accept all the terms of this license and intellectual property
(trademark/copyright) agreement. If you do not agree to abide by all
the terms of this agreement, you must cease using and return or
destroy all copies of Project Gutenberg™ electronic works in your
possession. If you paid a fee for obtaining a copy of or access to a
Project Gutenberg™ electronic work and you do not agree to be
bound by the terms of this agreement, you may obtain a refund
from the person or entity to whom you paid the fee as set forth in
paragraph 1.E.8.

1.B. “Project Gutenberg” is a registered trademark. It may only be

used on or associated in any way with an electronic work by people
who agree to be bound by the terms of this agreement. There are a
few things that you can do with most Project Gutenberg™ electronic
works even without complying with the full terms of this agreement.
See paragraph 1.C below. There are a lot of things you can do with
Project Gutenberg™ electronic works if you follow the terms of this
agreement and help preserve free future access to Project
Gutenberg™ electronic works. See paragraph 1.E below.
1.C. The Project Gutenberg Literary Archive Foundation (“the
Foundation” or PGLAF), owns a compilation copyright in the
collection of Project Gutenberg™ electronic works. Nearly all the
individual works in the collection are in the public domain in the
United States. If an individual work is unprotected by copyright law
in the United States and you are located in the United States, we do
not claim a right to prevent you from copying, distributing,
performing, displaying or creating derivative works based on the
work as long as all references to Project Gutenberg are removed. Of
course, we hope that you will support the Project Gutenberg™
mission of promoting free access to electronic works by freely
sharing Project Gutenberg™ works in compliance with the terms of
this agreement for keeping the Project Gutenberg™ name associated
with the work. You can easily comply with the terms of this
agreement by keeping this work in the same format with its attached
full Project Gutenberg™ License when you share it without charge
with others.

1.D. The copyright laws of the place where you are located also
govern what you can do with this work. Copyright laws in most
countries are in a constant state of change. If you are outside the
United States, check the laws of your country in addition to the
terms of this agreement before downloading, copying, displaying,
performing, distributing or creating derivative works based on this
work or any other Project Gutenberg™ work. The Foundation makes
no representations concerning the copyright status of any work in
any country other than the United States.

1.E. Unless you have removed all references to Project Gutenberg:

1.E.1. The following sentence, with active links to, or other

immediate access to, the full Project Gutenberg™ License must
appear prominently whenever any copy of a Project Gutenberg™
work (any work on which the phrase “Project Gutenberg” appears,
or with which the phrase “Project Gutenberg” is associated) is
accessed, displayed, performed, viewed, copied or distributed:
 This eBook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it away
or re-use it under the terms of the Project Gutenberg License
included with this eBook or online at www.gutenberg.org. If you
are not located in the United States, you will have to check the
laws of the country where you are located before using this
eBook.

1.E.2. If an individual Project Gutenberg™ electronic work is derived

from texts not protected by U.S. copyright law (does not contain a
notice indicating that it is posted with permission of the copyright
holder), the work can be copied and distributed to anyone in the
United States without paying any fees or charges. If you are
redistributing or providing access to a work with the phrase “Project
Gutenberg” associated with or appearing on the work, you must
comply either with the requirements of paragraphs 1.E.1 through
1.E.7 or obtain permission for the use of the work and the Project
Gutenberg™ trademark as set forth in paragraphs 1.E.8 or 1.E.9.

1.E.3. If an individual Project Gutenberg™ electronic work is posted

with the permission of the copyright holder, your use and distribution
must comply with both paragraphs 1.E.1 through 1.E.7 and any
additional terms imposed by the copyright holder. Additional terms
will be linked to the Project Gutenberg™ License for all works posted
with the permission of the copyright holder found at the beginning
of this work.

1.E.4. Do not unlink or detach or remove the full Project

Gutenberg™ License terms from this work, or any files containing a
part of this work or any other work associated with Project
Gutenberg™.

1.E.5. Do not copy, display, perform, distribute or redistribute this

electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1
with active links or immediate access to the full terms of the Project
Gutenberg™ License.

1.E.6. You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form,
including any word processing or hypertext form. However, if you
provide access to or distribute copies of a Project Gutenberg™ work
in a format other than “Plain Vanilla ASCII” or other format used in
the official version posted on the official Project Gutenberg™ website
(www.gutenberg.org), you must, at no additional cost, fee or
expense to the user, provide a copy, a means of exporting a copy, or
a means of obtaining a copy upon request, of the work in its original
“Plain Vanilla ASCII” or other form. Any alternate format must
include the full Project Gutenberg™ License as specified in
paragraph 1.E.1.

1.E.7. Do not charge a fee for access to, viewing, displaying,

performing, copying or distributing any Project Gutenberg™ works
unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8. You may charge a reasonable fee for copies of or providing

access to or distributing Project Gutenberg™ electronic works
provided that:

• You pay a royalty fee of 20% of the gross profits you derive
from the use of Project Gutenberg™ works calculated using the
method you already use to calculate your applicable taxes. The
fee is owed to the owner of the Project Gutenberg™ trademark,
but he has agreed to donate royalties under this paragraph to
the Project Gutenberg Literary Archive Foundation. Royalty
payments must be paid within 60 days following each date on
which you prepare (or are legally required to prepare) your
periodic tax returns. Royalty payments should be clearly marked
as such and sent to the Project Gutenberg Literary Archive
Foundation at the address specified in Section 4, “Information
 about donations to the Project Gutenberg Literary Archive
Foundation.”

• You provide a full refund of any money paid by a user who

notifies you in writing (or by e-mail) within 30 days of receipt
that s/he does not agree to the terms of the full Project
Gutenberg™ License. You must require such a user to return or
destroy all copies of the works possessed in a physical medium
and discontinue all use of and all access to other copies of
Project Gutenberg™ works.

• You provide, in accordance with paragraph 1.F.3, a full refund of

any money paid for a work or a replacement copy, if a defect in
the electronic work is discovered and reported to you within 90
days of receipt of the work.

• You comply with all other terms of this agreement for free
distribution of Project Gutenberg™ works.

1.E.9. If you wish to charge a fee or distribute a Project Gutenberg™

electronic work or group of works on different terms than are set
forth in this agreement, you must obtain permission in writing from
the Project Gutenberg Literary Archive Foundation, the manager of
the Project Gutenberg™ trademark. Contact the Foundation as set
forth in Section 3 below.

1.F.

1.F.1. Project Gutenberg volunteers and employees expend

considerable effort to identify, do copyright research on, transcribe
and proofread works not protected by U.S. copyright law in creating
the Project Gutenberg™ collection. Despite these efforts, Project
Gutenberg™ electronic works, and the medium on which they may
be stored, may contain “Defects,” such as, but not limited to,
incomplete, inaccurate or corrupt data, transcription errors, a
copyright or other intellectual property infringement, a defective or
damaged disk or other medium, a computer virus, or computer
codes that damage or cannot be read by your equipment.

1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for

the “Right of Replacement or Refund” described in paragraph 1.F.3,
the Project Gutenberg Literary Archive Foundation, the owner of the
Project Gutenberg™ trademark, and any other party distributing a
Project Gutenberg™ electronic work under this agreement, disclaim
all liability to you for damages, costs and expenses, including legal
fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR
NEGLIGENCE, STRICT LIABILITY, BREACH OF WARRANTY OR
BREACH OF CONTRACT EXCEPT THOSE PROVIDED IN PARAGRAPH
1.F.3. YOU AGREE THAT THE FOUNDATION, THE TRADEMARK
OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL
NOT BE LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT,
CONSEQUENTIAL, PUNITIVE OR INCIDENTAL DAMAGES EVEN IF
YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH DAMAGE.

1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you

discover a defect in this electronic work within 90 days of receiving
it, you can receive a refund of the money (if any) you paid for it by
sending a written explanation to the person you received the work
from. If you received the work on a physical medium, you must
return the medium with your written explanation. The person or
entity that provided you with the defective work may elect to provide
a replacement copy in lieu of a refund. If you received the work
electronically, the person or entity providing it to you may choose to
give you a second opportunity to receive the work electronically in
lieu of a refund. If the second copy is also defective, you may
demand a refund in writing without further opportunities to fix the
problem.

1.F.4. Except for the limited right of replacement or refund set forth
in paragraph 1.F.3, this work is provided to you ‘AS-IS’, WITH NO
OTHER WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED,
INCLUDING BUT NOT LIMITED TO WARRANTIES OF
MERCHANTABILITY OR FITNESS FOR ANY PURPOSE.

1.F.5. Some states do not allow disclaimers of certain implied

warranties or the exclusion or limitation of certain types of damages.
If any disclaimer or limitation set forth in this agreement violates the
law of the state applicable to this agreement, the agreement shall be
interpreted to make the maximum disclaimer or limitation permitted
by the applicable state law. The invalidity or unenforceability of any
provision of this agreement shall not void the remaining provisions.

1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation,

the trademark owner, any agent or employee of the Foundation,
anyone providing copies of Project Gutenberg™ electronic works in
accordance with this agreement, and any volunteers associated with
the production, promotion and distribution of Project Gutenberg™
electronic works, harmless from all liability, costs and expenses,
including legal fees, that arise directly or indirectly from any of the
following which you do or cause to occur: (a) distribution of this or
any Project Gutenberg™ work, (b) alteration, modification, or
additions or deletions to any Project Gutenberg™ work, and (c) any
Defect you cause.

Section 2. Information about the Mission

of Project Gutenberg™
Project Gutenberg™ is synonymous with the free distribution of
electronic works in formats readable by the widest variety of
computers including obsolete, old, middle-aged and new computers.
It exists because of the efforts of hundreds of volunteers and
donations from people in all walks of life.

Volunteers and financial support to provide volunteers with the

assistance they need are critical to reaching Project Gutenberg™’s
goals and ensuring that the Project Gutenberg™ collection will
remain freely available for generations to come. In 2001, the Project
Gutenberg Literary Archive Foundation was created to provide a
secure and permanent future for Project Gutenberg™ and future
generations. To learn more about the Project Gutenberg Literary
Archive Foundation and how your efforts and donations can help,
see Sections 3 and 4 and the Foundation information page at
www.gutenberg.org.

Section 3. Information about the Project

Gutenberg Literary Archive Foundation
The Project Gutenberg Literary Archive Foundation is a non-profit
501(c)(3) educational corporation organized under the laws of the
state of Mississippi and granted tax exempt status by the Internal
Revenue Service. The Foundation’s EIN or federal tax identification
number is 64-6221541. Contributions to the Project Gutenberg
Literary Archive Foundation are tax deductible to the full extent
permitted by U.S. federal laws and your state’s laws.

The Foundation’s business office is located at 809 North 1500 West,

Salt Lake City, UT 84116, (801) 596-1887. Email contact links and up
to date contact information can be found at the Foundation’s website
and official page at www.gutenberg.org/contact

Section 4. Information about Donations to

the Project Gutenberg Literary Archive
Foundation
Project Gutenberg™ depends upon and cannot survive without
widespread public support and donations to carry out its mission of
increasing the number of public domain and licensed works that can
be freely distributed in machine-readable form accessible by the
widest array of equipment including outdated equipment. Many
small donations ($1 to $5,000) are particularly important to
maintaining tax exempt status with the IRS.

The Foundation is committed to complying with the laws regulating

charities and charitable donations in all 50 states of the United
States. Compliance requirements are not uniform and it takes a
considerable effort, much paperwork and many fees to meet and
keep up with these requirements. We do not solicit donations in
locations where we have not received written confirmation of
compliance. To SEND DONATIONS or determine the status of
compliance for any particular state visit www.gutenberg.org/donate.

While we cannot and do not solicit contributions from states where

we have not met the solicitation requirements, we know of no
prohibition against accepting unsolicited donations from donors in
such states who approach us with offers to donate.

International donations are gratefully accepted, but we cannot make

any statements concerning tax treatment of donations received from
outside the United States. U.S. laws alone swamp our small staff.

Please check the Project Gutenberg web pages for current donation
methods and addresses. Donations are accepted in a number of
other ways including checks, online payments and credit card
donations. To donate, please visit: www.gutenberg.org/donate.

Section 5. General Information About

Project Gutenberg™ electronic works
Professor Michael S. Hart was the originator of the Project
Gutenberg™ concept of a library of electronic works that could be
freely shared with anyone. For forty years, he produced and
distributed Project Gutenberg™ eBooks with only a loose network of
volunteer support.
Project Gutenberg™ eBooks are often created from several printed
editions, all of which are confirmed as not protected by copyright in
the U.S. unless a copyright notice is included. Thus, we do not
necessarily keep eBooks in compliance with any particular paper
edition.

Most people start at our website which has the main PG search
facility: www.gutenberg.org.

This website includes information about Project Gutenberg™,

including how to make donations to the Project Gutenberg Literary
Archive Foundation, how to help produce our new eBooks, and how
to subscribe to our email newsletter to hear about new eBooks.
Welcome to Our Bookstore - The Ultimate Destination for Book Lovers
Are you passionate about books and eager to explore new worlds of
knowledge? At our website, we offer a vast collection of books that
cater to every interest and age group. From classic literature to
specialized publications, self-help books, and children’s stories, we
have it all! Each book is a gateway to new adventures, helping you
expand your knowledge and nourish your soul
Experience Convenient and Enjoyable Book Shopping Our website is more
than just an online bookstore—it’s a bridge connecting readers to the
timeless values of culture and wisdom. With a sleek and user-friendly
interface and a smart search system, you can find your favorite books
quickly and easily. Enjoy special promotions, fast home delivery, and
a seamless shopping experience that saves you time and enhances your
love for reading.
Let us accompany you on the journey of exploring knowledge and
personal growth!

ebookgate.com