Weed-Crop

Competition
A Review
Second Edition
Weed-Crop
Competition
A Review
Second Edition

Robert L. Zimdahl
Robert L. Zimdahl is Professor of Weed Science, Authorization to photocopy items for internal or per-
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Blackwell Publishing Ltd
9600 Garsington Road, Oxford OX4 2DQ, UK Zimdahl, Robert L.
Tel.: +44 (0)1865 776868 Weed-crop competition : a review / Robert L.
Zimdahl—2nd ed.
Blackwell Publishing Asia p. cm.
550 Swanston Street, Carlton, Victoria 3053, Australia Includes bibliographical references (p.).
Tel.: +61 (0)3 8359 1011 ISBN 0-8138-0279-2 (alk. paper)
1. Weeds. 2. Plant competition. 3. Weeds—
Bibliography. 4. Plant competition—Bibliography.
I. Title.
SB611.Z55 2004
632'.5—dc22
2003023656

The last digit is the print number: 9 8 7 6 5 4 3 2 1

 In order to penetrate ever further into their subjects, the host of specialists narrow
their fields and dig down deeper and deeper till they can’t see each other from hole
to hole. But the treasures their toil brings to light they place on the ground above.
A different kind of specialist should be sitting there, the only one still missing.
He would not go down any hole, but would stay on top and piece all the different
facts together.
—Thor Heyerdahl, Aku-Aku: The Secret of Easter Island
 Contents

Preface ix

1 Introduction: An Historical Perspective 1

2 Definition of Plant Competition 6
3 Competition in the Community 9
Plant Communities 9
Agricultural Communities 10
Levels of Competition 10
Density 13
Community Composition 14
Theories of Competition 16
4 Influence of Competition on the Plant 19
Density 19
Competitive Ability 19
General Principles 20
Competitive Success 22
5 The Effect of Weed Density 27
Alfalfa 28
Barley 28
Corn/Maize 32
Cotton 39
Oilseed Crops 46
Flax 46
Rapeseed/Canola 46
Safflower 48
Sunflower 48
Peanut/Groundnut 49
Potato 51
Rice 52
Sorghum 56
Soybean 57
Sugarbeet 75
Sugarcane 77
Vegetables 78
Bean 78
Lentil and Chickpea 80

vii
viii Contents

Onion 81
Pea 81
Pepper 82
Tomato 83
Other Vegetable Crops 86
Wheat 88
Other Small Grain Crops 103
Studies of Diverse Crops 104
Weed-Weed Interference 106
6 The Effect of Competition Duration 109
7 The Elements of Competition 131
The Role of Temperature 132
Competitive Interactions for Nutrients 133
Competitive Interactions for Light 136
Competitive Interactions for Water 139
Competition for Other Environmental Factors 141
8 Weed Management Using the Principles of Competition 146
Plant Arrangement in the Community 147
Monoculture Versus Polyculture 149
Tillage 150
Rotation or Crop Sequence 153
Shade 155
The Role of Crop Genotype 156
Fertility 159
The Importance of Weed Biology and Ecology 160
9 Methods Used to Study Weed-Crop Competition 167
10 Models and Modeling 173
Conceptual Models 174
Simulation (Analytical) Models 174
Mechanistic or Empirical Models 176
Time of Emergence 179
Leaf Area Models 179
Multispecies Competition 182
The Extrapolation Domain of Models 183
Decision-Aid Models 183
Spatial Distribution 185
The Effect of Variability on Decisions 186
Thresholds 187
Conclusion 190
11 Conclusion: The Complexity of Competition 197

Appendix 200
Index 211
 Preface
The primary impetus for the first edition of this
work twenty years ago came from my opinion,
formed from limited international experience in the
1970s, that many weed scientists in developing
countries did not receive and were not aware of cur-
rent weed science literature (see Zimdahl 1980).
They had limited or no access to journals common-
ly found in libraries of the developed world. Thus,
they were denied use of printed resources that help
develop an historical perspective. Often, they did not
know what was known. An historical perspective
combined with the stimulation of current research
sharpens the focus of research programs and facili-
tates their justification to administrators and funding
agencies. Lack of access to the literature can narrow
one’s perspective and usually impedes development
of good weed science programs.
Because no comprehensive review of weed-crop
competition had been published and because
approval had been given for the project by the Inter-
national Plant Protection Center at Oregon State
University, there was additional motivation for the
first edition.
The literature review for the present book began
in mid-2001 in the library of the International Rice
Research Institute (IRRI), Los Banos, Laguna,
Philippines. The review was completed and writing
began in late 2002. When the review began, I was a
Fulbright Scholar in the Department of Agronomy
of the University of the Philippines at Los Banos,
and Dr. James Hill, chair of IRRI’s Department of
Crop, Soil, and Water Sciences, graciously offered a
courtesy appointment and access to IRRI facilities.
The Philippines, a place where weeds grow abun-
dantly, was an appropriate location to begin to think
again about what may be the central hypothesis of
ix
weed science: Weeds compete with crops and
reduce crop yield and quality.
This hypothesis is rarely stated in scientific
papers about weeds because it has dominated the
thinking in weed science for so long that it is
axiomatic. After all, if it were proven to be false and
if it were discovered that crops tolerated weeds, the
world would not need weed scientists. There would
be no problems with weed-crop competition. How-
ever, the first edition of this book, published in 1980,
showed that weed-crop competition is real and its
effects had been studied in many ways, in many
crops, for many years. The hypothesis that weeds
negatively affect crop yield and quality has been
tested and verified; it is accepted.
However, weed science, similar to most disci-
plines, continues to test its central hypothesis. Weed
scientists have been productively engaged in what
Kuhn (1970) calls normal science—“the activity in
which most scientists inevitably spend almost all
their time.” It is, in Kuhn’s view, “predicated on the
assumption that the scientific community knows
what the world is like.” It is a “strenuous attempt to
force nature into the conceptual boxes supplied by
professional education.” Thus, the weed science
community has continued to test its central hypoth-
esis about how weeds negatively affect crops. Weed
scientists are moving, albeit slowly, from the normal
science that repetitively asks what happens,
although this review establishes that these experi-
ments are still done, to the more difficult but more
important and more scientifically demanding ques-
tion of why does what is observed occur.
The first edition of this book was a report of what
had been done by whom. It included articles direct-
ly related to weed-crop competition published prior
x Preface
to June 1978. This second edition is an attempt to
summarize the literature about what is known about
what happens and to explore current understanding
of why. A goal is to urge a decrease of effort direct-
ed toward answering the first question and an
increase of effort on the second. In spite of criticism
of what has been done, I hasten to add that I have
been continually impressed with the quality of the
work and by the people who have done it. I have
been most impressed by many of the papers and
reviews mentioned here that are superlative work
done by capable people whose scientific knowledge
and skills often seem to extend beyond my analyti-
cal and review ability. I am humbled by what my
colleagues have done.
Unless warranted, this second edition will not
reconsider but will include some of the manuscripts
used in the first edition. To this end, I begin this
review at the end of the first edition and go forward.
Older material is included for historical reasons and
to make certain points. The book’s focus is interfer-
ence in the narrow sense of crop-weed competition.
The abundant recent literature on weed biology and
weed ecology is not included unless such studies
directly address competition. There was no attempt
to include any of the literature on allelopathy, which
has been summarized by others (Inderjit et al. 1995,
1999; Putnam and Tang 1986; Rice 1974, 1979,
1983; Thompson 1985). It is my limitation, but in
most cases, the review includes only literature pub-
lished in English with emphasis on American and
European journals of weed science. There are
exceptions, but, in general, this review does not
include papers published in the proceedings or
research progress reports of U.S. regional (e.g.,
northeastern, north central, southern, or western)
societies of weed science and by other regional
weed conferences (e.g., Asian-Pacific Weed Science
Society, Canadian Weed Science Society). This is
the case because the review emphasizes papers that
have passed peer review and been published in ref-
ereed journals. Second, many regional publications
were not readily available to me. Finally, papers that
emphasized herbicides or other weed management
techniques have not been reviewed. Readers will
note that much of what has been included seems
repetitious. Roget’s thesaurus helps, but not much,
when one wants to say that someone or a paper
showed, discovered, revealed, noted, or found. The
ways to say what was discovered are limited, espe-
cially when so much of the work included is similar
in design and result. I assure the reader that I recog-
nized the risk and suffered while writing from repe-
tition. The work assembled here is a resource, and I
hope one result will be that a lot more work to
explain what happens when weeds interfere with
crop growth will not be done.
Authors resist and editors insist on uniformity and
a limited set of notations and measurement systems.
The current convention of using only metric units
was tempting. However, readers who elected to con-
sult a particular paper would need to convert back to
the original units. Therefore, the units from original
papers were used without conversion to metric. A
short conversion table has been included as appen-
dix table A.4.
All weeds are cited by the common or scientific
name (if no common name has been accepted by the
Weed Science Society of America). Equivalent sci-
entific and common names, accepted by the Weed
Science Society of America, are included in appen-
dix table A.2, which lists them in alphabetical order
by common name, and in appendix table A.3, which
lists them in alphabetical order by scientific name.
The scientific name of each crop is included in
appendix table A.1.
Most papers selected for inclusion specifically
discuss weed-crop competition. Others provide
background information. Most literature concerning
crop-crop interactions has been omitted as has that
dealing with environmental conditions that stress
crops (e.g., low water, high temperature) and
increase their susceptibility to weed competition.
The second edition follows the general outline of
the first edition. A chapter on modeling and a more
detailed chapter on methods have been added.
LITERATURE CITED
Inderjit, K. M. M. Dakshini, and C. L. Foy, ed. 1999.
Principles and practices in plant ecology:
Allelochemical interactions. Boca Raton, FL: CRC
Press.
Inderjit, K. M. M. Dakshini, and F. A. Einhellig, ed.
1995. Allelopathy: Organisms, processes, and
applications. Amer. Chem. Soc. Symp. Series No.
268. Washington, DC.
Kuhn, T. S. 1970. The structure of scientific
revolutions. 2d ed. Chicago: University of Chicago
Press.
Putnam, A. R., and Chung-Shih Tang, ed. 1986. The
science of allelopathy. New York: John Wiley and
Sons.
 Preface xi

Rice, E. L. 1974. Allelopathy. New York: Academic
Press.
———. 1979. Allelopathy—An update. Bot. Rev.
45:15–109.
———. 1983. Pest control with nature’s chemicals:
Allelochemicals and pheromones in gardening and
agriculture. Norman: Oklahoma University Press.
Thompson, A. C., ed. 1985. The chemistry of
allelopathy: Biochemical interactions among plants.
Amer. Chem. Soc. Symp. Series No. 268.
Washington, DC.
Zimdahl, R. L. 1980. Weed-crop competition: A
review. Corvallis, OR: Int. Plant Prot. Center,
Oregon State University.