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A Review
Second Edition
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.
Zimdahl2nd 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 competitionBibliography.
I. Title.
SB611.Z55 2004

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 cant 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

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

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

The primary impetus for the first edition of this weed science: Weeds compete with crops and
work twenty years ago came from my opinion, reduce crop yield and quality.
formed from limited international experience in the This hypothesis is rarely stated in scientific
1970s, that many weed scientists in developing papers about weeds because it has dominated the
countries did not receive and were not aware of cur- thinking in weed science for so long that it is
rent weed science literature (see Zimdahl 1980). axiomatic. After all, if it were proven to be false and
They had limited or no access to journals common- if it were discovered that crops tolerated weeds, the
ly found in libraries of the developed world. Thus, world would not need weed scientists. There would
they were denied use of printed resources that help be no problems with weed-crop competition. How-
develop an historical perspective. Often, they did not ever, the first edition of this book, published in 1980,
know what was known. An historical perspective showed that weed-crop competition is real and its
combined with the stimulation of current research effects had been studied in many ways, in many
sharpens the focus of research programs and facili- crops, for many years. The hypothesis that weeds
tates their justification to administrators and funding negatively affect crop yield and quality has been
agencies. Lack of access to the literature can narrow tested and verified; it is accepted.
ones perspective and usually impedes development However, weed science, similar to most disci-
of good weed science programs. plines, continues to test its central hypothesis. Weed
Because no comprehensive review of weed-crop scientists have been productively engaged in what
competition had been published and because Kuhn (1970) calls normal sciencethe activity in
approval had been given for the project by the Inter- which most scientists inevitably spend almost all
national Plant Protection Center at Oregon State their time. It is, in Kuhns view, predicated on the
University, there was additional motivation for the assumption that the scientific community knows
first edition. what the world is like. It is a strenuous attempt to
The literature review for the present book began force nature into the conceptual boxes supplied by
in mid-2001 in the library of the International Rice professional education. Thus, the weed science
Research Institute (IRRI), Los Banos, Laguna, community has continued to test its central hypoth-
Philippines. The review was completed and writing esis about how weeds negatively affect crops. Weed
began in late 2002. When the review began, I was a scientists are moving, albeit slowly, from the normal
Fulbright Scholar in the Department of Agronomy science that repetitively asks what happens,
of the University of the Philippines at Los Banos, although this review establishes that these experi-
and Dr. James Hill, chair of IRRIs Department of ments are still done, to the more difficult but more
Crop, Soil, and Water Sciences, graciously offered a important and more scientifically demanding ques-
courtesy appointment and access to IRRI facilities. tion of why does what is observed occur.
The Philippines, a place where weeds grow abun- The first edition of this book was a report of what
dantly, was an appropriate location to begin to think had been done by whom. It included articles direct-
again about what may be the central hypothesis of ly related to weed-crop competition published prior

x Preface

to June 1978. This second edition is an attempt to in design and result. I assure the reader that I recog-
summarize the literature about what is known about nized the risk and suffered while writing from repe-
what happens and to explore current understanding tition. The work assembled here is a resource, and I
of why. A goal is to urge a decrease of effort direct- hope one result will be that a lot more work to
ed toward answering the first question and an explain what happens when weeds interfere with
increase of effort on the second. In spite of criticism crop growth will not be done.
of what has been done, I hasten to add that I have Authors resist and editors insist on uniformity and
been continually impressed with the quality of the a limited set of notations and measurement systems.
work and by the people who have done it. I have The current convention of using only metric units
been most impressed by many of the papers and was tempting. However, readers who elected to con-
reviews mentioned here that are superlative work sult a particular paper would need to convert back to
done by capable people whose scientific knowledge the original units. Therefore, the units from original
and skills often seem to extend beyond my analyti- papers were used without conversion to metric. A
cal and review ability. I am humbled by what my short conversion table has been included as appen-
colleagues have done. dix table A.4.
Unless warranted, this second edition will not All weeds are cited by the common or scientific
reconsider but will include some of the manuscripts name (if no common name has been accepted by the
used in the first edition. To this end, I begin this Weed Science Society of America). Equivalent sci-
review at the end of the first edition and go forward. entific and common names, accepted by the Weed
Older material is included for historical reasons and Science Society of America, are included in appen-
to make certain points. The books focus is interfer- dix table A.2, which lists them in alphabetical order
ence in the narrow sense of crop-weed competition. by common name, and in appendix table A.3, which
The abundant recent literature on weed biology and lists them in alphabetical order by scientific name.
weed ecology is not included unless such studies The scientific name of each crop is included in
directly address competition. There was no attempt appendix table A.1.
to include any of the literature on allelopathy, which Most papers selected for inclusion specifically
has been summarized by others (Inderjit et al. 1995, discuss weed-crop competition. Others provide
1999; Putnam and Tang 1986; Rice 1974, 1979, background information. Most literature concerning
1983; Thompson 1985). It is my limitation, but in crop-crop interactions has been omitted as has that
most cases, the review includes only literature pub- dealing with environmental conditions that stress
lished in English with emphasis on American and crops (e.g., low water, high temperature) and
European journals of weed science. There are increase their susceptibility to weed competition.
exceptions, but, in general, this review does not The second edition follows the general outline of
include papers published in the proceedings or the first edition. A chapter on modeling and a more
research progress reports of U.S. regional (e.g., detailed chapter on methods have been added.
northeastern, north central, southern, or western)
societies of weed science and by other regional LITERATURE CITED
weed conferences (e.g., Asian-Pacific Weed Science
Inderjit, K. M. M. Dakshini, and C. L. Foy, ed. 1999.
Society, Canadian Weed Science Society). This is
Principles and practices in plant ecology:
the case because the review emphasizes papers that
Allelochemical interactions. Boca Raton, FL: CRC
have passed peer review and been published in ref-
ereed journals. Second, many regional publications Inderjit, K. M. M. Dakshini, and F. A. Einhellig, ed.
were not readily available to me. Finally, papers that 1995. Allelopathy: Organisms, processes, and
emphasized herbicides or other weed management applications. Amer. Chem. Soc. Symp. Series No.
techniques have not been reviewed. Readers will 268. Washington, DC.
note that much of what has been included seems Kuhn, T. S. 1970. The structure of scientific
repetitious. Rogets thesaurus helps, but not much, revolutions. 2d ed. Chicago: University of Chicago
when one wants to say that someone or a paper Press.
showed, discovered, revealed, noted, or found. The Putnam, A. R., and Chung-Shih Tang, ed. 1986. The
ways to say what was discovered are limited, espe- science of allelopathy. New York: John Wiley and
cially when so much of the work included is similar Sons.
Preface xi

Rice, E. L. 1974. Allelopathy. New York: Academic Thompson, A. C., ed. 1985. The chemistry of
Press. allelopathy: Biochemical interactions among plants.
. 1979. AllelopathyAn update. Bot. Rev. Amer. Chem. Soc. Symp. Series No. 268.
45:15109. Washington, DC.
. 1983. Pest control with natures chemicals: Zimdahl, R. L. 1980. Weed-crop competition: A
Allelochemicals and pheromones in gardening and review. Corvallis, OR: Int. Plant Prot. Center,
agriculture. Norman: Oklahoma University Press. Oregon State University.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

An Historical Perspective

Two of the earliest references on the effects of weed stances left by preceding crops. One of the first stud-
competition appear in ancient writings of the Bible: ies of plant competition was by Sachs (1860), who
Cursed is the ground for thy sake; in sorrow shalt attempted to relate soil mass to plant yield. Nageli in
thou eat of it all the days of thy life; thorns also and 1865 (cited in Clements et al. 1929) broadened the
thistles shall it bring forth to thee; and thou shalt eat significance of competition in the plant community,
the herb of the field (Genesis 3:1718). Another pointing out that it furnished a solution to the prob-
passage, the parable of the sower, notes that, some lems involved with the presence of lime in soil.
fell among thorns; and the thorns sprang up, and Malthus (1798) was concerned primarily with
choked them (Matthew 13:7). growth of the human population and the consequent
It is correct to suggest that competition among poverty and misery he saw in Liverpool, England.
plants precedes recorded history and that it was rec- He proposed that the power of population was infi-
ognized long before a defined term was assigned. nitely greater than the earths ability to produce
Competition among plants in natural communities is food. The Malthusian apocalypse, when the human
common, but it is not a universal phenomenon population is greater than the ability of the earth to
(Goldberg 1990). However, in agricultural plant produce food, has been avoided because of develop-
communities, weed-crop competition, with a few ments in food production technology. Competition
exceptions (e.g., living mulches, companion crop- for food, if the Malthusian apocalypse comes true,
ping), seems to be a natural, undesirable, ubiquitous, still concerns many. Malthus said, The cause to
and inevitable phenomenon. which I allude is the constant tendency in all ani-
Competition is a predictable response of grouping mated life to increase beyond the nourishment pre-
living organisms into communities. Clements et al. pared for it. The Malthusian hypothesis has three
(1929, p. 3) provided an early history of the litera- major points:
ture, which they claimed was not extensive. The
1. Population is necessarily limited by the
book by Clements et al. (1929), although perhaps
means of subsistence.
not extensive, provides an accurate historical per-
2. Population infinitely increases when the
spective on the early development of the study of
means of subsistence increase unless it is
plant competition. Competition was recognized and
controlled by powerful checks.
reported by Petrus de Crescentiis in 1305 when he
3. These checks that suppress the superior
directed that trees in a forest community be cut first
power of population and keep population and
where they were too thick. The significance of com-
the means of subsistence in balance are
petition in the plant kingdom was perhaps first per-
resolvable by moral restraint, or population
ceived by Decandolle (1820) who described plant
will be controlled by the four horsemen of
competition and stated that all species of a region
the apocalypse: war, famine, pestilence, and
and all plants of a given place are in a state of war
with respect to each other. He derived a theory of
antagonism between phanerogams, and a theory of Malthuss views are claimed by some to have
crop rotation based on the idea that succeeding been inspiration for Darwins (1859) principle of
species should be those not inhibited by toxic sub- natural selection. Darwin (1859) derived the concept

2 Chapter 1

of competition (a struggle for existence) in natural ments of plant competitionlight, nutrients, and
evolution and proposed it was ubiquitous and waterare mentioned repeatedly in the literature.
omnipresent. In reviewing Darwins exposition of Portions of this review will focus on each, but we
competition in the Origin of the Species, one can must recognize that plants exist in environments
easily overlook the fact that he regarded competition where all elements of competition are active, and
as only one component of the struggle for existence, separation of the elements, while interesting, proba-
but possibly the most important one. bly does not, and perhaps cannot, reflect the real
Nageli (cited in Clements et al. 1929) sought to world of inter- and intraspecific competition.
give mathematical form to the suppression of plants Plant competition has been studied from two major
by their competitors. He concluded that the number perspectives. Ecologists have studied competition to
of species in an area was determined by the average understand diversity and change in plant communities
life period and the average annual growth increment. and patterns of succession in plant communities. The
In 1892, Macmillan (cited in Clements et al. 1929) goal has been to develop principles for management of
considered the significance of competition between ecosystems. Agronomists, weed ecologists, and weed
communities as well as within them. He was among scientists (the difference between the last two cate-
the first to express the view that there are certain gories is small) have been most interested in competi-
points of resemblance in the competition for food tion between weeds, the unwanted species, and crops.
that takes place between similar individuals and Studies have emphasized weed and crop density, rela-
causes the weaker to be more or less suppressed. tive time of weed and crop emergence, and the use of
Clements et al. (1929) stated, herbicides and other management techniques to mini-
mize weed competition in crops. This review empha-
Competition is a question of the reaction of the plant sizes the work of weed scientists but does not exclude
upon the physical factors that encompass it and of the complementary work by plant ecologists. The review
effect of these modified factors upon adjacent plants.
In the exact sense, two plants do not compete as long
intentionally excludes the many good studies on weed
as the water-content, the nutrients, the heat and light control.
are in excess of the needs of both. The moment, how- Kropff and Walter (2000) noted that the introduc-
ever, that the roots of one enter the area from which tion of selective herbicides has been one of the main
the other draws its water supply, or the foliage of one
factors enabling intensification of agriculture in
begins to overshadow the leaves of the other, the reac-
tion of the former modifies unfavorably the factors con- developed countries. During its first three decades,
trolling the latter, and competition is at once initiated. the journal Weed Research published mainly papers
(pp. 1011) related to herbicides. As reported by Kropff and Wal-
ter (2000), most papers published now are on weed
Haldane (1932) noted, Fitness depends in a quite biology and ecology (32 in 1998 versus 17 in 1975)
complicated way on the environment. In order to test with fewer on herbicides (6 in 1998 versus 32 in
fitness in the Darwinian sense it would be necessary 1975). The work reported in Weed Science has
to grow the plants in competition. Thus, we must shown the same shift in emphasis (fig. 1.1). Thirty
conclude that a complete analysis of plant competi- years ago (1973), 82 percent of the papers published
tion must involve plants in a community and their in Weed Science dealt with some aspect of herbi-
communal relations, plus individual plant-growth cides, and 18 percent dealt with some aspect of weed
patterns. The growth of plants in isolation can pro- biology/ecology. Herbicide-related studies have
vide useful information but only when it is com- steadily declined to only 37 percent of the papers in
bined with community studies. 2003, whereas 63 percent emphasize some aspect of
Brenchley (1917) studied several weeds in culti- weed biology/ecology. Figure 1.1 shows the steady
vated crops and observed that some were generally decrease of herbicide papers and the steady increase
found in association with certain crops and others of those on weed biology/ecology with the latter
were common among all cultivated crops. She exceeding the former in 1999.
hypothesized that one of the foremost factors deter- Kropff and Walter (2000) regard the shift in
mining a particular weed species abundance or emphasis from herbicides to weed biology and ecol-
scarcity was its ability to withstand competition. ogy as desirable. They note Mortensen et al.s
The aboveground struggle for light, she stated, was (2000) report of significant application of weed
as important as the underground competition for biology research to solving practical weed manage-
nutrients and moisture. These three primary ele- ment problems.
Introduction: An Historical Perspective 3

Figure 1.1. A 30-year account of papers published on herbicides and on weed biology and
ecology in Weed Science.. Herbicide articles included all articles that focused on herbicides in any
way (e.g., physiology, mycoherbicides, herbicide resistance). Biology/ecology articles were select-
ed only if they did not mention herbicides. Statistical and modeling papers were excluded.

Barbieri and Kropff (2002), two years later, chemical use. This is achievable, in their view,
reported a further shift in emphasis. They said that through integration of weed science with other dis-
in weed biology we are moving from prediction of ciplines (e.g., ecology and environmental science),
the time of weed emergence to quantification of the and that integration in Barbieri and Kropffs (2002)
process, taking particularly into account the vari- view may be the beginning of the loss of identity of
ability of weather conditions. They suggest that weed science as a stand-alone discipline.
there is an increasing number of people studying It is not an objective of this book to engage in the
the interactions between weeds and other biota. legitimate debate about whether weed science has
Barbieri and Kropff (2002) also suggest that crop- ever achieved standing as a discipline with its own
weed competition studies are increasingly applied to university departments, curriculum, and degrees.
scenarios in which biological and cultural weed con- However, perhaps weed scientists ought to or will be
trol methods play a major role. This review does not compelled by the direction of their science to begin
deny their observation but neither does it offer vig- to consider their future carefully and the benefits
orous confirmation. Barbieri and Kropff (2002) pro- and losses of the desirable integration proposed by
pose that if this trend toward system-thinking is Barbieri and Kropff (2002).
real, it should be seen as a positive achievement for To prepare this book, all issues of the worlds
weed scientists. It is positive because it will pro- three principal weed science journals: Weed Science,
duce research results that give farmers and con- Weed Technology, and Weed Research from 1980 to
sumers what they really want from weed science mid-2002 were reviewed for articles that dealt with
research. It is apparent that Barbieri and Kropff weed-crop competition. It is almost surely true that
(2002) think that what farmers and consumers real- some important papers were missed. The error is
ly want is improved accuracy and precision of weed- mine and I apologize to those whose papers were
management recommendations and reduced missed and for the incomplete report that may be the
4 Chapter 1

result. Earlier volumes were reviewed for the first weed scientists as the detrimental effect of weeds on
edition. It is interesting to note how many articles in crops, and they have been expressed as a reduction
each issue of each journal dealt specifically with of crop yield. This legitimate weed-science approach
weed-crop competition (table 1.1) and to compare is contrasted with the ecological approach that
those data with those in figure 1.1. emphasizes the mechanism(s) of competition in
After this introduction and a brief historical per- plant communities and the structure of communities.
spective on plant competition, chapter 2 focuses on The chapters provide evidence for the desirable con-
the definition of competition as it is used in the vergence of the two approaches.
weed-crop literature, which recapitulates the mater- Chapter 5, the largest part of this edition,
ial that appeared in the first edition of this book. reviews the many reports of the result of competi-
Chapters 3 and 4 cover the role of competition in tion between several weeds in several crops. It pro-
the community and the influence of competition on vides abundant support for the hypothesis that
individual plants. These chapters are brief not increasing weed density decreases crop yield.
because they are unimportant but because these are Chapter 5 is arranged alphabetically by crop, and
not the areas that weed-science research has empha- the literature for each crop is placed at the end of
sized. The effects of competition in the crop-weed that crops section rather than at the end of the
community have nearly always been measured by whole chapter.

Table 1.1. Number of Articles on Weed-Crop Competition in Three Major Weed Science
Journals, 1979 to 2002

Year Weed Research Weed Science Weed Technology

1979 3 11
1980 4 18
1981 1 16
1982 2 16
1983 5 10
1984 7 31
1985 2 20
1986 3 31
1987 3 34 4
1988 5 26 6
1989 5 24 3
1990 4 17 7
1991 6 17 14
1992 8 28 8
1993 7 21 4
1994 8 18 9
1995 8 21 5
1996 7 36 4
1997 5 17 3
1998 8 11 1
1999 9 17 0
2000 5 6 7
2001 2 16 8
Mid-2002 8 9 5
Note: Weed Technology was first published in 1987.
Introduction: An Historical Perspective 5

Chapter 6 summarizes some of the literature in LITERATURE CITED

chapter 5 that describes the effect of competition
Barbieri, P., and M. J. Kropff. 2002. Twelfth European
duration. Chapter 5 emphasizes the effects of weed
Weed Res. Soc. Symp.: What is the trend in weed
density (how many), whereas chapter 6 emphasizes research? Eur. Weed Res. Soc. Newsletter, No. 81.
the variation introduced by the duration (how long) of Brenchley, W. E. 1917. The effect of weeds upon
competition. Chapter 6 also reexamines the critical- cereal crops. New Phytologist 16:5376.
period concept. There is an abundance of data that Clements, F. E., J. E. Weaver, and H. C. Hanson.
purport to define a critical period for weed manage- 1929. Plant competitionan analysis of
ment in several different crops, but the critical period community function. Publ. No. 398. Carnegie
seems peripheral to most weed management deci- Institute, Washington, DC.
sions. It is clear that the period varies with the place Darwin, C. 1859. On the origin of species by means of
where the work was done, with the particular season, natural selection0r the preservation of favoured
with each weed-crop combination, and with the rela- races in the struggle for life. Albemarle St.,
tive time of emergence of the crop and weed(s). The London: J. Marray. A facsimile of the first edition,
data on the critical period for weed control are inter- 1967. New York: Atheneum.
esting but may be useful only where the work was DeCandolle, A. P. 1820. Essai lmentaire de
done and not generalizable. gographie botanique (cited in Clements et al.
Chapter 7 attempts to separate, as the literature 1929).
frequently does, the things (light, nutrients, water) Goldberg, D. H. 1990. Components of resource
that plants compete for, which plants that must inte- competition in plant communities. In Perspectives
grate cannot separate. on plant competition, ed. J. B. Grace and D.
Tilman, 2749. San Diego, CA: Academic Press.
Chapter 8 asks if all the work that has been done
Haldane, J. B. S. 1966. The causes of evolution.
has made a difference. Has weed management
Ithaca, NY: Cornell University Press (First pub.
improved because of all that has been done to estab-
1932, Longmans, Green and Co., London).
lish that weeds reduce crop yield? The answer,
Kropff, M. J., and H. Walter. 2000. EWRS and the
sadly, seems to be that while the work is not useless, challenges for weed research at the start of the new
the evidence that it has made a major difference in millennium. Weed Res. 40:710.
weed management techniques is weak. Malthus, T. R. 1798. An essay on the principle of
Chapter 9 explores the methods used to study population as it affects the future improvement of
weed-crop competition. These were mentioned in society, with remarks on the speculation of Mr.
the first edition, but the importance of the experi- Godwin, M. Condorcet, and other writers. London,
mental method to eventual interpretation of data was England, printed for J. Johnson in St. Pauls
not emphasized in the first edition. church-yard.
Chapter 10 is new because so little had been done Mortensen, D. A., L. Bastianns, and M. Sattin. 2000.
on models before 1979. Now much has been done, The role of ecology in the development of weed
and models and modeling are important to develop- management systems: An outlook. Weed Res.
ment of improved weed management methods. 40:4962.
Chapter 11 attempts to conclude the results of Ngeli, C. 1874. Verdrngung der Pflanzenformen
more than 650 research papers. There are three durch ihre Mitbewerber. Ib 11:109164, 3:205260.
major conclusions. The first is that there is no ques- 1881.
tion that what I have called the central hypothesis of Sachs, J. 1860. Bericht ber physiologische Thtigkeit
weed science has been affirmed: Weeds compete an der Versuchstation in Thrandt. II. Wurzelstudien.
with crops and reduce crop yield and quality. The Landw. Versuchs. 2:131.
work that has been done affirms this for numerous
weeds in all of the important crops that have been
studied. The second conclusion is that weed science
will benefit from closer integration with plant ecol-
ogy and greater emphasis on study and understand-
ing of the coexistence of plants. The final conclusion
is that modeling has become an important aspect of
modern weed management systems and is likely to
become more important.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

Definition of Plant Competition

Where there is so much of competition and around them and the modified environment in
uncertainty you must expect self interest will turn influences the growth of the constituent
govern. plants.
3. Bleasdale (1960): Two plants are in
Jeremy Collier, Essays on Moral Subjects,
competition with each other when the growth
of either one or both of them is reduced or
their form modified as compared with their
The Oxford English Dictionary defines competition
growth or form in isolation.
as the action of endeavouring to gain what another
4. Milne (1961): Competition is the endeavor
endeavours to gain at the same time; the striving of
of two (or more) animals to gain the measure
two or more for the same object; rivalry. To compete
each wants from the supply of a thing when
comes from the Latin word competere, which means
that supply is not sufficient for both (or all).
to ask or sue for the same thing another does. Despite
5. Birch (1955): Competition occurs when a
a precise etymology and concise definition, the
number of animals (of the same or of
meaning of the term competition is not precise in the
different species) utilize common resources
plant science literature. Milne (1961) proposed that
the supply of which is short; or if the
confusion resulted from (1) a misunderstanding of
resources are not in short supply, competition
Darwins original usage, (2) neglect of the etymolo-
occurs when the animals seeking that
gy of the word, and (3) the mixing of competition
resource nevertheless harm one or the other
with results. Milne, who worked with animals, found
in the process.
wide disagreement among definitions.
6. Clements et al. (1929): Competition involves
Bunting (1960) thought competition had different
the reaction of a plant to the physical factors
shades of meaning for the agronomist and the plant
that encompass it and of the effect of these
physiologist. In his view, physiologists think of
modified factors upon adjacent plants. For
competition as being for something, usually nutri-
Clements et al., competition is a purely
ents, water, or light. Agronomists and weed scien-
physical process.
tists, while agreeing, add that competition also
exists between plants (Donald 1963) or parts of the In the exact sense, two plantsno matter how
same plant. The many definitions were reviewed by close, do not compete with each other so long as
Milne (1961). the water content, the nutrient material, the light
and heat are in excess of the needs of
1. Mather (1961): Competition implies the both....Competition occurs when each of two or
presence of one individual as an effective part more organisms seeks the measure they want of
any particular factor or things and when the imme-
of the others environment and a similarity of diate supply of the factor or things is below the
need or activities so their impact on each combined demand of the organisms.
other is prospectively detrimental.
2. Aspinall and Milthorpe (1959): The This definition makes competition different from
interaction between plants and environment. the broader term interference, which includes com-
Plants during growth modify the environment petition, and allelopathy.

Definition of Plant Competition 7

Harper (1960, 1961) decided that many defini- only two broad categories of mechanisms: direct
tions proved excessively cumbersome and, in his interference and indirect exploitation of shared
work, adopted and has argued for use of the inclu- resources. In the first case, two plants compete
sive term, interference, which has been advocated directly for a resource both require. In the second
by Muller (1969). As mentioned in the preface, dis- case, exploitative competition is an indirect action
cussion of allelopathy has not been included in this because of the requirement for a shared resource.
review. Allelopathy is distinguished from competi- Connell (1990) notes that if one wants to distinguish
tion because it depends on adding a chemical com- the types of competition (he cites four possibilities),
pound to the environment whereas competition knowledge of the mechanisms is required. He agrees
involves the removal or reduction of an essential with Tilman (1987) that few studies of competition
factor from the environment. Competition, as weed by ecologists have demonstrated the mechanisms
scientists understand it and use the term, is a strug- underlying the observed interactions, and therefore
gle between a crop and a weed for a resource (e.g., it is quite possible that many studies that purport to
light, water, or nutrients) that is in short supply demonstrate competition may illustrate apparent,
Grace and Tilman (1990) point out that defini- not real, competition. The lack of study of the mech-
tions of competition range from the narrow to the anisms is certainly true for studies of weed-crop
general, from operational to philosophical, and from competition performed by weed scientists. However,
phenomenological to mechanistic. The range of as Radosevich and Roush (1990) point out (see
definitions, in their view, has caused confusion and chapter 9), while this is true, it should not be viewed
continues to cloud discussion of the substance of as a devastating critique because the objectives of
competition. The many definitions notwithstand- the studies performed by weed scientists have been
ing, weed scientists concerned with weed-crop com- distinctly different from the objectives of ecologists.
petition discover, with pleasure, the two major Perhaps the simplest distinction between the worthy
points of plant competition outlined by Clements et objectives of the two groups is that the former is
al. (1929), followed by an inclusive definition. The interested in what happens and the latter in why.
principles are: One often reads of competition for space, and
while this occurs in the animal kingdom, it is not
1. Competition is keenest when individuals are
usually the case with plants. Rivalry for space may
most similar and make the same demands on
occur with sugarbeet or carrot when two roots actu-
the habitat and adjust themselves less readily
ally touch or become intertwined. Generally, in
to their mutual interactions.
plant competition, the phrase implies competition
2. The closeness of competition between plants
for the resources space containsnutrients, water,
of different species varies directly with their
or lightrather than for the space itself.
likeness in vegetation or habitat form.
Further evidence that the association of two or
Dissimilarity tends to eliminate competition
more plant species does not always result in compe-
and preserve the advantage of the superior
tition can be found in the symbiotic association of
form. These broad concepts seem to fit with
legumes and grasses. Mather (1961) discussed this
the weed scientists perception of weed-crop
aspect of plant relationships and Donald (1963) pro-
competition and are compatible with the
vided another example: the germinating seeds of
work that has been done.
subterranean clover. The dormancy found in some
Donald (1963) combined the definitions of Milne varieties of this species for many weeks after har-
(1961) for the animal world and Clements et al. vesting may be broken by exposure to an atmos-
(1929) for plants into a concise statement: Compe- phere containing 0.5 percent carbon dioxide. If one
tition occurs when each of two or more organisms seed in a dormant group germinates in a normal
seeks the measure it wants of any particular factor or atmosphere, it will provide enough carbon dioxide
thing and when the immediate supply of the factor to initiate germination of the rest. Work on living
or thing is below the combined demand of the mulches and companion crops also points to the fact
organisms. that not all plant combinations are competitive or
Connell (1990) defined competition as simply a harmful.
reciprocal negative interaction between two organ- Donald (1963) also mentioned that competition
isms. Connell (1990) points out that ecologists tra- cannot be assumed simply because a factor is in
ditionally restrict the term to instances involving short supply. If all plants in a community are
8 Chapter 2

exposed to insufficiency while the environment of Clements, F. E., J. E. Weaver, and H. C. Hanson.
each is independent of its neighbors, there can be no 1929. Plant competitionan analysis of
competition. He used the example of poor oxygen community function. Publ. No. 398. Washington,
supply delaying germination and growth of wheat DC: Carnegie Institute.
seedlings in very wet, poorly structured soils. How- Connell, J. H. 1990. Apparent versus real
ever, the circumstance is exceptional and competi- competition in plants. In Perspectives on plant
tion soon occurs. competition, ed. J. B. Grace and D. Tilman, 926.
Plants cannot be considered to compete for heat San Diego, CA: Academic Press.
because heat is not present in finite amounts, Donald, C. M. 1963. Competition among crop and
pasture plants. Advances in Agron. 15:1118.
although heat can affect the process (see chapter 7).
Academic Press, NY.
Competition for carbon dioxide may occur but prob-
Grace, J., and D. Tilman. 1990. Perspectives on plant
ably only under extremely crowded conditions. Most
competition: Some introductory remarks. In
of the factors for which there is competition are Perspectives on plant competition, ed. J. B. Grace
found as a pool from which supplies are drawn, and D. Tilman, 3, 4. San Diego, CA: Academic
according to Donald (1963). This concept can be Press.
easily visualized for water and nutrients but not for Harper, J. L. 1960. Factors controlling plant numbers.
light. Light must be intercepted when available or it In The biology of weeds, ed. J. L. Harper, 119132.
is lost forever. Thus, foliar height and breadth will Oxford, UK: Blackwell Sci. Publ.
determine a plants effectiveness as a competitor for . 1961. Approaches to the study of plant
light. One can refer to Clements et al. (1929) to fur- competition. Symposium Soc. for Exp. Biol.
ther elucidate the concept of a pool and its usage: It 15:139.
is evident that practically all the advantages or Mather, K. 1961. Competition and cooperation. In
weapons of competing species are epitomized in two Mechanisms in biological competition, Soc. for
wordsamount and rate. Greater storage in seed or Exp. Biol. Symp. XV. Academic Press, NY.
rootstock, more rapid and complete germination, 15:264281.
earlier start, more rapid growth of roots and shoots, Milne, A. 1961. Definition of competition among
taller and more branching stems, deeper and more animals. In Mechanisms in biological competition,
spreading roots, more tillers, larger leaves, and more Soc. for Exp. Biol. Symp. XV. Academic Press, NY.
numerous flowers are all of the essence of success. 15:4061.
Thus, nothing succeeds like success. Muller, C. H. 1969. Allelopathy: A factor in
Competitive ability has been proposed as a genet- ecological process. Vegetatio 18:348357.
Oxford English Dictionary. 1971 ed., s.v.
ic character controlled by polygenes but not associ-
ated with morphological characters such as height,
Radosevich, S. R., and M. L. Roush. 1990. The role
growing habit, and vigor of growth (Sakai 1961).
of competition in agriculture. In Perspectives on
The heritability of the trait, if it exists, appears to be plant competition, ed. J. B. Grace and D. Tilman,
very low and the outcome of competition surely 341363. San Diego, CA: Academic Press.
must vary not only with intensity, but also with the Sakai, K. 1961. Competitive ability in plants: Its
environment in which it occurs. inheritance and some related problems. In
LITERATURE CITED Mechanisms in biological competition, Soc. for
Exp. Biol. Symp. XV. Academic Press, NY. 15:
Aspinall, D., and F. L. Milthorpe. 1959. An analysis 245263.
of competition between barley and white persicaria. Tilman, D. 1987. The importance of the mechanisms
Ann. Appl. Bio. 47: 156172. of interspecific competition. Am. Nat. 129:769774.
Birch, L. C. 1955. Selection in Drosophila
pseudoobscura in relation to crowding. Evolution
Bleasdale, J. K. A. 1960. Studies on plant
competition. In The biology of weeds, ed. J. L.
Harper, 133142. Oxford, UK: Blackwell Sci. Publ.
Bunting, A. H. 1960. Some reflections on the ecology
of weeds. In The biology of weeds, ed. J. L. Harper,
1126. Oxford, UK: Blackwell Sci. Publ.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

Competition in the Community

The effects of competition in the crop-weed com- has not been a problem for the weed-science commu-
munity have nearly always been measured by weed nity because it has been assumed to occur, unequivo-
scientists as the effect of the weed or weeds on the cally, albeit with the necessary semantic combination
crop, and the effect is commonly expressed as a of competition and allelopathy under the general term
reduction in crop yield. The importance of competi- interference. Both competitive and noncompetitive
tion as a determinant of the structure of natural plant processes will (if both occur in a field) strongly influ-
communities is well accepted and it has been evalu- ence plant growth in a multispecies community (Hall
ated in several ways (Connell 1990). In Connells 1974). Connell (1990) states the evidence for compe-
view, the first task is to demonstrate unequivocally tition (e.g., crop yield reduction) can often result from
its occurrence in nature, yet this has often proved to other types of plant interactions, which are frequently
be difficult. Goldberg (1990) concurs that competi- overlooked. These are illustrated in figure 3.1 (Con-
tion is a common although not ubiquitous phenom- nell 1990).
Research for this book strongly suggests weed sci-
entists regard competition as ubiquitous in agricultur- As weed scientists expand the search for mech-
al plant communities. Demonstration of its occurrence anisms of competition and study the structure of

Figure 3.1. Types of real and apparent competition among plants. Solid lines are direct interac-
tions, and dashed lines are indirect. An arrowhead shows a positive effect on that species, and a
circle shows a negative effect. In case 4, apparent competition is between P1 and P3. (From Con-
nell 1990; reprinted with permission)

10 Chapter 3

crop-weed communities, the difference between have one or more of the following characteristics:
direct and indirect competition and apparent compe-
1. Different nutritional requirements, as illus-
tition will become more important. An example of
trated by legumes and grasses coexisting in
the kind of work that weed science needs more of is
pasture and hay fields.
the work on community assembly by Booth and
2. Different causes of mortality, observed in
Swanton (2002). Community assembly, a rare term
pastures where animals selectively graze.
in weed science, is a branch of ecology that exam-
3. Different sensitivity to toxins, including alle-
ines how a community is assembled over time and
lochemicals and herbicides.
what paths (trajectories) the members and the entire
4. A different time demand for growth factors.
community follow over time.
Many plants require the same growth factors
The trajectories are determined by biotic (competi-
to succeed, but they do not demand them at
tion) and nonbiotic factors (environment). Booth and
the same time. This may be the most com-
Swanton call these filters, each of which acts at multi-
mon reason for coexistence.
ple scales. Environmental filters act to remove or limit
species that lack specific traits. Booth and Swanton A more complete discussion of coexistence is pre-
present the basic ecological theory of community sented by Grime (1979, pp. 157177).
assembly and propose how it can be applied to weed-
science research to predict how crop-weed communi-
ties change in response to what they call imposed In agricultural fields, orderly, continuous, natural
filters such as tillage and crop rotation. Their work, if ecological succession does not occur, but change,
followed, may lead weed science toward fundamental due to human manipulation, does. Agricultural
theories of competition and away from continued fields that are not in a permanent crop such as an
emphasis on what happens. They acknowledge that a orchard or have a semipermanent perennial crop
community assembly approach is the opposite of the (e.g., alfalfa, where disturbance is still frequent for
current approach to weed management, which is to harvest and the land is rotated to another crop per-
look at weeds as a series of individual problems and to haps as often as every three years) are regularly and
study the biology of each species in an attempt to iden- intentionally disturbed and lack a natural (undis-
tify weak links in their life cycle. Management tech- turbed) plant community. Environmental change is a
niques are then developed to address the identified driving force in natural plant succession but annual
weak links at specific sites under specific conditions. crop agriculture strives to modify and control the
Booth and Swanton (2002) recognize the utility of environment through tillage, fertilization, pest
the approach but caution that it does little to broad- (weed, insect, and disease) control, and irrigation.
en understanding of why weeds occur where they do Because of regular disturbance and other cultural
or how they interact in communities. The approach practices, the spatial and temporal variability of agri-
leads to solution of the weed problem addressed and cultural environments is reduced compared to natural
creation of a new weed problem that the solution did plant communities (Radosevich and Roush 1990).
not (could not) address. It is the kind of solution crit- Dominance of the planted crop species characterizes
icized by Berry (1981) as one that leads to a rami- agricultural plant communities that also have a few
fying series of new problems. Berry (1981) (rarely only one) weed species that occur in cropped
advocated, as Booth and Swanton (2002) do, solu- fields. Their removal (control) creates open niches
tions that cause a ramifying series of solutions. into which another weedy species will move, but per-
Understanding how communities are assembled and haps not immediately. Therefore, weed management,
function should lead to a series of solutions. especially successful weed management, is a never-
The community, in which usually a single crop ending process (Zimdahl 1999). One could argue that
and one-to-many weed species exist, is important to the best weed management techniques may therefore
weed management and the study of competition be those that achieve less than 100 percent control
because it is the organizational level at which and do not open niches that allow new introductions
changes occur. Changes can occur within a species to succeed and necessitate further weed control.
by mutation and ecotype development or by replace-
ment of one species with another (Zimdahl 1999,
pp. 130131). There are at least four reasons why The three primary processes that control the level of
two or more species can coexist in a place. They can competition from a weed complex in a crop are
Competition in the Community 11

described by Radosevich and Roush (1990) and conflicting truism that competition within plant
have been recognized by many others. Time of communities is intense and should therefore have
emergence of the crop and weed often determine the important evolutionary consequences, but that plant
outcome of interference. When the crop emerges species coexist with apparently little differentiation
prior to the weeds, it often wins the competitive bat- that permits interaction avoidance.
tle; the reverse is also true. Firbank and Watkinson Monoculture rarely occurs in natural environ-
(1985) reported that emergence time and local ments because communal life is favored. Even what
crowding accounted for as much as 50 percent of the appears to the casual observer to be a monocultural
variation in performance of individual corn cockle plant community (e.g., a large field of corn) teems
plants. A second factor is growth ability and envi- with other species (bacteria and fungi on or in
ronment (Radosevich and Roush 1990). These are plants, weeds, insects, soil microorganisms, etc.).
related because growth is surely affected by envi- Nature does not recognize the human categories of
ronment, but they can also be treated separately domesticated plant, or such things as inalienable
because growth of some species (e.g., green foxtail rights. In natural environments, most living organ-
and field bindweed) will always be distinctly differ- isms are engaged in relentless competition for
ent, independent of the environment in which they resources with peers as well as with many other
are growing together. organisms. That competition, which we know as
Clearly, plant growth rate has a strong influence part of the process of evolution, in a real sense, con-
on competitive ability. Species that grow tall rapidly serves past evolutionary achievements by protecting
and gain greater ground cover (shading) or spread features that assured success.
rapidly laterally will have a competitive advantage In the ecological sense, competition and natural
over those that do each thing but more slowly. selection, as parts of the evolutionary process, gen-
Equally clearly, environments that favor rapid erate and conserve what is valuable for survival;
growth will favor the species with greater competi- they create better fitness1. Plants do not escape the
tive ability. Finally, Radosevich and Roush (1990) struggle for existence and the competitive process
mention the important but often neglected role(s) that creates better fitness. As pointed out above,
played by processes other than competition, such as competition, a part of evolution, is common but is
herbivory, density-dependent mortality, predation, not ubiquitous in all natural communities (Goldberg
senescence, and allelopathy. 1990). It is common and assumed to be ubiquitous
Aarssen (1989) stated that plant species coexist in agricultural communities where its results have
by avoiding competitive exclusion. He agreed with been studied by weed scientists and others interest-
Sakai (1961) when he proposed that, at the species ed in weed-crop competition. Goldbergs (1990)
level, continuing selection results from genetically thought asks for separation of the effect of competi-
based differences in competitive abilities in local tion from the plants response to competition. The
neighborhoods (fig. 3.2). Aarssen reported on multi- effect could be on the abundance of one competitor
generation experiments with timothy and common and the response could be on the abundance of the
groundsel to show that competitive ability may other or on its yield. Weed-science research has
change as a consequence of selection. He explored tended to focus on the response, especially of the
the evolutionary consequences of such selection in a crop, with less attention to the effect that causes the
community of several species. The resulting hypoth- response.
esis suggested that competitive exclusion is avoided Brenchley (1917) emphasized the omnipresence
at the population level because no population con- of competition as a vital factor in the agricultural
tains even one genotype that is competitively supe- plant community when she said, It is impossible to
rior to all other genotypes in a coexisting sow a crop without the certainty that other plants
population. Aarssens (1989) hypothesis assumes will appear. Pavlychenko and Harrington (1934)
that competitive ability is intransitive (not capable showed that competition exerts a powerful natural
of transition to another species). If the intransitive force in the agricultural plant community tending
characteristic extends across several species, then toward limitation or extinction of weaker competi-
the most competitive genotypes are just as likely to tors. They found that, within the community, each
belong to one species as to another. Aarssen (1989) weed and each plant differed greatly in competitive
concludes that the hypothesis of competitive com- ability and that all weeds suffered greatly from com-
bining ability helps to explain what he calls the petition with crop plants.
12 Chapter 3

Figure 3.2.
A proposed
among attributes
of competitive
ability in plants.
Primary traits are
those that have
the most
important role in
determining the
competitive ability
of neighbors.
Some primary
traits are
determined by
secondary traits
and some
secondary traits
are determined
by tertiary traits.
(From Aarssen
1989; reprinted
with permission)
Competition in the Community 13

DENSITY (Mann and Barnes 1947) depended on which

became established first in the community and bar-
Donald (1963) began a discussion of density in the
ley seeding rate. If the weed established during the
community with an examination of the relationship
previous year, a thin stand of barley could reduce
of density to total yield of dry matter, the biological
barley yield up to 100 percent. Competition of bar-
yield of various crops (as distinct, for example, from
ley with common chickweed varied slightly in that
the yield of grain). He pointed out that studies to
the weed reduced the yield of barley, but the oppo-
determine optimum sowing rate rarely include a suf-
site relation was not true, primarily because of more
ficiently wide range of densities to permit definition
rapid development of common chickweed (Mann
of the relationship of density and yield. The data
and Barnes 1950). The work by Mann and Barnes is
from Donalds experiment (1951) on intraspecific
similar to many other studies that show the effect of
competition among annual pasture plants indicate
weeds on a crop in terms of crop yield.
the relation of yield of dry matter to density at zero
Topham and Lawson (1982) asked if crop inter-
days (weight of seed embryo), 131 days, and 181
ference changes the weed flora over time. Using
days in subterranean clover grown with adequate
diversity indices, they showed that despite increas-
moisture and nutrients. At planting, there was a lin-
ing substantial competitive pressure from vining
ear relation between density and yield. Competition
peas on dry matter accumulation by up to eight
for light developed in dense populations soon after
weeds, weed species composition did not change
germination and thereafter became operative in pop-
with time. Competition from peas suppressed weed
ulations of lower and lower density. Competitive
growth and reduced the number of species identified
effects stopped growth at highest densities. Because
on successive sampling dates. However, there was
of extreme growth rate reduction late in the season
no selection pressure and species evenness was
and concurrent high growth rate in sparse stands,
unchanged. In the view of Topham and Lawson
sparse stands tended to approach the more dense
(1982), the use of diversity indices is ecologically
stand in final yield. The final data showed that yield
justified and will aid weed management decisions.
of dry matter is constant from moderate to high den-
A comment by Salisbury (1942) summarizes:
sities. The original linear relationship of density to
Below a certain specific density the increased yield
yield of dry matter was replaced by a curve in which
of the individual fails to compensate for the dimin-
yield rose sharply with increasing density to a max-
ished population. On the other hand, above a certain
imum, which was constant for all higher densities.
density, the individual becomes so depauperate
This work also stresses that the determination of
through competition, that the augmented population
optimum density for an early harvest is more diffi-
fails to compensate for the low yield of the individ-
cult than at maturity. For an early harvest, the
uals. Although Salisbury was speaking of an indi-
greater the density the greater the yield because the
vidual in a monoculture, his reasoning can be
earlier harvest will come at a time when intercrop
extrapolated to the community.
competition is less intense with consequent lower
The literature on competition leads to the problem
yield depression.
of determining if a given unit of soil will produce a
The early work of Aspinall and Milthorpe (1959)
fixed increment of growth and yield with the pre-
presents a similar relationship by analyzing compe-
vailing environment, or if the competitive influence
tition between barley and pale smartweed where a
in annual crops intervenes to reduce total yield in
constancy of final yield of dry matter per unit area at
favor of the yield of one component of a population.
moderate to high densities was measured.
The experiment by Aspinall and Milthorpe (1959)
Mann and Barnes (1945, 1947, 1949, 1950,
has been cited in this regard. Robinson and Dunham
1952), in several carefully conducted competition
(1954) found that soybeans produce normal yields,
experiments, demonstrated that yield of crop and
and sometimes more, when forage companion crops
weeds tended to a maximum with a definite density
were interseeded in soybean rows. As corn was
of plants per volume of soil. However, effects
intercropped with mung bean and the level of weed
between plants were inconsistent. Corn spurry and
control was reduced, the relative advantage of inter-
scentless chamomile limited the growth of barley
cropping increased so corns productivity was 75
and were, in turn, limited by barley. The same held
percent greater than in a monoculture (Bantilan et al.
true for clovers (Mann and Barnes 1952) and the
1974). The response of weeds was correlated with
grass redtop (Mann and Barnes 1949), in competi-
light interception ability.
tion with barley. The effect of German velvetgrass
14 Chapter 3

Mann and Barnes (1945) showed that with a con- patchiness, generalizations that are useful when
stant amount of weediness from either of two weeds, making management decisions cannot be made with
an increasing density of barley plants diminished the reference to species and environment, which vary
injurious effects of the weeds. The combined weight among experiments. Natural environments favor life
of barley and weeds was rarely as great as barley in a diverse community for plants. While pasture
alone in a weed-free plot. The research team stated and hay crops are usually seeded as mixtures, most
that with a constant density of barley and a variable developed country agriculture relies on monocul-
density of weeds, the total weight of the above- ture.
ground portion of barley and weeds was almost con- The question of possible yield advantages for
stant, independent of the number of weed plants of mixed-culture communities persists. In this regard,
either species. without going into great detail, the work of Wes
Moolani and Slife (1960) found that dry weight of Jackson (1980, 1987; and Vitek and Jackson 1996)
weeds and corn combined was equal to the weight of at the Land Institute (of Salina, Kansas) on high
weed-free corn. However, with soybeans, the crop seed yielding, perennial polycultures is worth noting
plus weeds equaled one and one-half times the yield and learning from. Jackson emphasizes the model of
of the weed-free crop. In another experiment with the prairie, a place that sponsors its own fertility and
corn and soybeans, Knake and Slife (1962) found pest management as a new, feasible, and proper par-
that increases in dry matter of giant foxtail were pro- adigm for the practice of agriculture. Interested
portional to decreases in dry matter of the crop. The readers are referred to the Land Institutes Land
combined yield did not vary significantly. Similar Report, which is published three times annually.
results were reported by Shadbolt and Holm (1948) Donald (1963) observed that native pasture com-
working with vegetables. munities commonly develop great complexity with
Staniforth often mentioned total yield in his work several layers of each species. He asked, Can such
with soybeans in Iowa, but the results were incon- a community structure exploit the environment to a
sistent. In two experiments (Staniforth 1958; Stani- maximum? If light-tolerant species will grow
forth and Weber 1956), yield of soybeans alone was beneath the canopy and if roots with varying degrees
almost equal to soybeans plus weeds. In other work of dispersal will better exploit available moisture
(Weber and Staniforth 1957), yield of soybeans and and nutrients, the answer may be yes. Plants inte-
weeds was slightly higher than when soybeans were grate all the variables in any environment. There-
grown alone. fore, an important agricultural and weed
Allison et al. (1958) present a possible mechanis- management question becomes: can two species fix
tic explanation. In a discussion of the relationship more carbon when growing in association than
between evapotranspiration losses and yield, they when either species grows alone? Are there yield
found a direct and high degree of correlation and weed management objectives that can be
between evapotranspiration and the dry weight pro- achieved through greater community diversity?
duced by aboveground parts. This was true regard- Clements et al. (1929) described competition for
less of crop rotation or fertility level. two factors: water and light. The beginning of com-
petition is due to reaction, when the plants are so
spaced that the reaction of one affects the response
The weight of evidence presented here suggests that of the other by limiting it. The initial advantage thus
few generalizations can be made about the constan- gained is increased by cumulation, since even a
cy of community composition or yield. Farmers and slight increase in the amount of energy, as raw mate-
weed scientists have known for some time that rial, is followed by corresponding growth, and this
weeds are not uniformly distributed across a field, by further gain in response and reaction. A larger,
that is, the communitys composition is highly vari- deeper, or more active root system enables one plant
able across a field. Weed management techniques to secure an amount of the chresard (available
have generally ignored this fact and managers have water), and the immediate reaction is to reduce the
chosen control methods that have been applied uni- amount obtainable to the other. The stem and leaves
formly to entire fields. The value of knowledge of the former grow in size and number, and thus
about patchiness and overall weed distribution will require more water; the roots respond by augment-
be to aid postemergence weed management deci- ing the absorbing surface to supply the demand and
sions (Wiles et al. 1992). Other than the existence of automatically reduce the water content still further.
Competition in the Community 15

At the same time, the correlated growth of stems and Studies of competition between associated
leaves is producing a reaction on light by absorp- species, other than forage crops, are rare, and per-
tion, leaving less energy available for the leaves of haps justifiably so because other crops are rarely
the competitor beneath it, while increasing the grown in the same way. Hanson et al. (1961) and
amount of food for the further growth of absorbing Hinson and Hanson (1962) found that the advantage
roots, taller stems, and overshading leaves. gained by one of a competing pair of soybean geno-
The view of Clements et al. (1929) strongly sug- types equaled the loss sustained by the other. They
gests that when two species grow together, one will considered competition between soybean genotypes
be suppressed while the other will dominate. This to be additive. Recent work with rice cultivars in
has been the dominant view of weed science. China has shown that a mixture of two cultivars in a
Ahlgren and Aamodt (1939), in contradiction, sug- single field outyields either grown alone.
gested that when some common mesophytic plants Stringfield (1959) observed that when two corn
are grown in pairs, the yield per plant of both species hybrids were grown in association, no marked
in the mixture may be less than the yield per plant in advantage or disadvantage in productivity accrued
each of the corresponding pure cultures. They tested to the mixture compared with the average of the
their hypothesis but did not substantiate it with a contributing hybrids when grown separately. The
mixture of redtop and Kentucky bluegrass. results were constant whether the members of a
Several experiments (Aberg et al. 1943; Donald given hybrid pair were alike or widely different. The
1963; Erdmann and Harrison 1947; Roberts and increase in yield of one was balanced by a decrease
Olsen 1942) with forage or grass species reveal in yield of the other. Two corn genotypes of differ-
extensive support for Donalds (1963) analysis of ent height were grown in association by Pendleton
possible results when two species grow together. To and Seif (1962). Alternate rows of a 106 inch tall
summarize: and a 72 inch tall variety were planted. In direct con-
trast to Stringfields (1959) results, the mixture
1. The yield of the mixture will usually be less
yielded 7 bu A-1 less than the mean of the two pure
than that of the higher-yielding pure culture.
cultures. The authors pointed out that considerable
2. The yield of the mixture will usually be
shading of the dwarf by the taller corn occurred, but
greater than that of the lower-yielding pure
there was only very ineffectual shading of the lower
leaves of the taller by the dwarf.
3. The yield of the mixture may be greater or
Overall, the limited data available indicate rela-
less than the mean yield of the two pure cul-
tively little gain from mixing species, but other
advantages obtained in certain environments, eco-
4. There is no substantial evidence that two
nomic situations, or crop rotations should not be dis-
species can exploit the environment better
missed. Harper and Gajic (1961) emphasized that
than one.
knowledge of factors controlling population in the
In other work, Donald (1958) indicated that com- plant community also determines the extent of
petition for two factors leads to multiple interactions understanding of the reasons for one species suc-
between two groups of effects and thus greatly ceeding at the expense of another. The same infor-
intensifies the effects of either factor operating mation helps explain why a diversity of plant
alone. Aggressor species showed a negative interac- species may cohabit in a relatively stable communi-
tion between the effect of two modes of competition ty without one becoming dominant while the popu-
(i.e., light and nutrient, or light and water). Yields lation of the other declines. It is commonly accepted
dropped slightly due to competition for either factor that two species scarcely ever occupy similar niches,
alone, but when competition for both factors operat- but displace each other so each takes possession of
ed, yields approached levels obtained in the absence certain resources, which gives it a competitive
of competition. The aggressor competed more effec- advantage. This view has been called the competi-
tively when both means of competition were avail- tive exclusion principle.
able to it. The effect of the two modes of The process of natural selection leads toward eco-
competition on the suppressed species showed a logical differentiation of competing species and
positive interaction. Yield depression, under compe- therefore promotes stability of ecosystems even
tition for both factors, greatly exceeded the sum of though competition is an unavoidable consequence
the effects of competition for the separate factors. of communities. Two plants or two competing
16 Chapter 3

species can compete in at least two different ways. acterized by species whose population can deplete
One species can rapidly deplete a resource required resources to a low level at some equilibrium state.
by both, or a species can continue to grow at Grimes theory is based on established and identifi-
depleted resource levels (Goldberg 1990). The able plant traits, whereas Tilmans theory is based
interaction between the effect on (competition for) on traits found in the population (Grace 1990).
resources and the response to resource depletion Grace (1990) suggests that the two theories are actu-
must also be considered (Goldberg 1990). A given ally complementary when the differences in the def-
plant may respond in both ways dependent on the inition of competition are considered. The
level of resources available. The responses could differences then become subtle, although not unim-
be positively or negatively correlated (Goldberg portant, in Graces (1990) view. Grime defines
1990). In some cases, plants that are able to survive competition as the capacity to capture resources
at reduced resource levels compete not by taking whereas Tilman defines it as a negative relationship
possession of certain resources but by gaining a between the abundances of competing species that
competitive advantage by being able to survive involves both capture and tolerance to low resource
with reduced resources; by consuming less. It is levels. Grace (1990) argues that Grimes definition
similar to a social principlethose who do best in is not operational, which is to say it does not con-
a time of declining resources may be those who can form to accepted definitions of plant competition.
make do with the least rather than those who must Grace (1990) says the problem with Tilmans theory
consume the most. is that it is consistent with conventional definitions,
The competitive ability of a species can be mea- but the operational definition is such that competi-
sured by how strongly it suppresses other individu- tion is the only factor leading to dominance (regard-
als (the net competitive effect) or by how little it less of disturbance rate or nonresource conditions).
responds to the presence of others (the net competi- The debate over the proper definition and mech-
tive response) (Goldberg 1990). These different but anism of competition in plant communities has not
related responses to competition can be measured by been resolved. The evidence from this review is
a plants effect on resources (how much is used how that weed-science research has not been opera-
quickly) or by its response to deficiency. In the first tionally affected by the debate. It will behoove the
case, a plant competes by consuming more rapidly weed-science community to pay attention to the
or in greater abundance than its neighbors. In the debate and make appropriate changes in research
second case, the plant responds to deficiency or to techniques and objectives. Changes or shifts over
greater consumption by a neighbor by increasing time of the weed species present in a disturbed
resource uptake, by decreasing resource loss, or by (cropped) community are a secondary effect of
increasing the efficiency of use of resources already weed management and may be independent of the
obtained, that is, reducing the requirement for addi- means. Intensive cropping systems give rise to
tional resources (Goldberg 1990). weed communities that are products of cropping
patterns and weed management systems rather than
a result of only natural competition and succes-
The difference in how plants deal with resources sion (Harwood et al. 1974). The fact that weed-crop
(compete or decline to compete directly) that is in competition takes place in such disturbed commu-
the definition of competitive ability is well charac- nities demands special techniques for study and
terized by the theories of competition proposed by analysis of results. It would be best if weed-science
Grime (1977, 1987) and Tilman (1982, 1985, 1988) research techniques were developed and employed
and discussed by Grace (1990). Grime (1977) in full consideration of the work of plant ecologists
thought competitive ability was directly related to in natural plant communities. That there is wisdom
traits that a plant possessed that permitted maxi- and value in ecological approaches to weed man-
mization of resource capture by individuals in a pop- agement is not a new plea and has been advocated
ulation. Consistent with the theory, Grime (1977) by others (Liebman and Dyck 1993).
posited that species that are stress tolerant (low
resource users) will dominate in nonsuccessional NOTE
communities, even if resources have been used by
other species (Goldberg 1990). In contrast, Tilman 1. I am indebted to my colleague Dr. Holmes
(1982) proposed that competitive ability was char- Rolston, III, of the Colorado State University
Department of Philosophy for this insight.
Competition in the Community 17

LITERATURE CITED plant competition, ed. J. B. Grace and D. Tilman,

2749. San Diego, CA: Academic Press.
Aarssen, L. W. 1989. Competitive ability and species
Grace, J. B. 1990. On the relationship between plant
coexistence: A plants-eye view. Oikos 56:386401.
traits and competitive ability. In Perspectives on
Aberg, E., I. J. Johnson, and C. P. Wilsie. 1943. Asso-
plant competition, ed. J. B. Grace and D. Tilman,
ciation between species of grasses and legumes.
5165. San Diego, CA: Academic Press.
Agron. J. 35: 357369.
Grime, J. P. 1977. Evidence for the existence of three
Ahlgren, H. L., and O. S. Aamodt. 1939. Harmful root
primary strategies in plants and its relevance to eco-
interactions as a possible explanation for effects
logical and evolutionary theory. Am. Nat.
noted between various species of grasses and
legumes. Agron. J. 31:982985.
. 1979. Plant strategies and vegetation
Allison, F. E., F. E. Roller, and W. A. Raney. 1958.
processes. Chichester, UK: J. Wiley & Sons.
Relationship between evapotranspiration and yields
. 1987. Dominant and subordinate components
of crops grown in lysimeters receiving natural rain-
of plant communities: Implications for succession,
fall. Agron. J. 50: 506511.
stability and diversity. In Colonization, succession
Aspinall, D., and F. L. Milthorpe. 1959. An analysis
and stability, ed. A. J. Gray, M. J. Crawley, and P.
of competition between barley and white persicaria.
J. Edwards, 413428. Oxford, UK: Blackwell Sci-
Ann. Appl. Biol. 47:156172.
Bantilan, R. T., M. C. Palada, and R. R. Harwood. ence Pubs.
1974. Integrated weed management: I, Key factors Hall, R. L. 1974. Analysis of the nature of interfer-
affecting crop-weed balance. Philippine Weed Sci. ence between plants of different species. I. Con-
Bul. 1:1436 cepts and extension of the de Wit analysis to
Berry, W. 1981. The gift of good land: Further essays examine effects. Aust. J. Agric. Res. 25:739747.
cultural and agricultural. San Francisco, CA: North Hanson, W. D., C. A. Brim, and K. Hinson. 1961.
Point Press. Design and analysis of competition studies with an
Booth, B. D., and C. J. Swanton. 2002. Assembly the- application to field plot competition in the soybean.
ory applied to weed communities. Weed Sci. Crop Sci. 1: 255258.
50:213. Harper, J. L., and D. Gajic. 1961. Experimental stud-
Brenchley, W. E. 1917. The effect of weeds upon ies of the mortality and plasticity of a weed. Weed
cereal crops. New Phytologist 16: 5376. Res. 1:91104.
Clements, F. E., J. E. Weaver, and H. C. Hanson. Harwood, R. R., and R. T. Bantilan. 1974. Integrated
1929. Plant competition b: An analysis of commu- weed management. II. Shifts in composition of the
nity function. Publ. No. 398. Washington, DC: weed community in intensive cropping systems.
Carnegie Inst. Phil. Weed Sci. Bull. 1:3759.
Connell, J. H. 1990. Apparent versus real competi- Hinson, K., and W. D. Hanson. 1962. Competition
tion in plants. In Perspectives on plant competition, studies in soybeans: Crop Sci. 2:117123.
ed. J. B. Grace and D. Tilman, 926. San Diego, Jackson, W. 1980. New roots for agriculture. San
CA: Academic Press. Francisco, CA: Friends of the Earth.
Donald, C. M. 1951. Competition among pasture . 1987. Altars of unhewn stone: Science and
plants. I. Intraspecific competition among annual the earth. San Francisco, CA: North Point Press.
pasture plants. Aust. J. Agr. Res. 2:355376. Knake, E. L., and F. W. Slife. 1962. Competition of
. 1958. The interaction of competition for light Setaria faberii with corn and soybeans. Weeds
and nutrients. Aust. J. Agric. Res. 9:421435. 10:2629.
. 1963. Competition among crop and pasture Liebman, M., and E. Dyck. 1993. Weed management:
plants. Adv. Agron. 15:118. A need to develop ecological approaches. Ecologi-
Erdmann, M. H., and C. M. Harrison. 1947. The cal Appl. 3:3941.
influence of domestic ryegrass and redtop upon the Mann, H. H., and T. W. Barnes. 1945. The competi-
growth of Kentucky bluegrass and Chewings fes- tion between barley and certain weeds under con-
cue in lawn and turf mixtures. Agron. J. trolled conditions. I. Competition with Spergula
39:682689. arvensis Linn. and Matricaria inodora Linn. Ann.
Firbank, L. G., and A. R. Watkinson. 1985. On the Appl. Biol. 32:1522.
analysis of competition within two species mixtures . 1947. The competition between barley and
of plants. J. Appl. Ecol. 22:503517. certain weeds under controlled conditions. II. Com-
Goldberg, D. H. 1990. Components of resource com- petition with Holcus mollis. Ann. Appl. Biol.
petition in plant communities. In Perspectives on 34:252266.
. 1949. The competition between barley and
18 Chapter 3

certain weeds under controlled conditions. III. Shadbolt, C. A., and L. G. Holm. 1948. A quantitative
Competition with Agrostis gigantea. Ann. Appl. study of the competition of weeds with vegetable
Biol. 39:273281. crops. Proc. North Cent. Weed Cont. Conf. 5:94.
. 1950. The competition between barley and Staniforth, D. W. 1958. Soybean-foxtail competition
certain weeds under controlled conditions. IV. under varying soil moisture conditions. Agron. J.
Competition with Stellaria media. Ann. Appl. Biol. 50:1315.
37:139148. Staniforth, D. W., and C. R. Weber. 1956. Effects of
. 1952. The competition between barley and annual weeds on the growth and yield of soybeans.
certain weeds under controlled conditions. V. Com- Agron. J. 48:467471.
petition with clover considered as a weed. Ann. Stringfield, G. H. 1959. Performance of corn hybrids
Appl. Biol. 39:111119. in mixtures. Agron. J. 51:472473.
Moolani, M. K., and F. W. Slife. 1960. The competi- Tilman, D. 1982. Resource competition and communi-
tive effects of various intensities of pigweed on the ty structure. Monographs in Population Biology.
development of corn and soybeans. Proc. North Princeton, NJ: Princeton University Press.
Cent. Weed Cont. Conf. 16:2627. . 1985. The resource-ratio hypothesis of plant
Pavlychenko, T. K., and J. B. Harrington. 1934. Com- succession. Am. Nat. 125:827852.
petitive efficiency of weeds and cereal crops. Cana- . 1988. Plant strategies and the dynamics of
dian J. Res. 10:7794. and structure of plant communities. Monographs in
Pendleton, J. W., and R. D. Seif. 1962. Role of height Population Biology. Princeton, NJ: Princeton Uni-
in competition. Crop Sci. 2:154156. versity Press.
Radosevich, S. R., and M. L. Roush. 1990. The role Topham, P. B., and H. M. Lawson. 1982. Measure-
of competition in agriculture. In Perspectives on ment of weed species diversity in crop/weed com-
plant competition, ed. J. B. Grace and D. Tilman, petition studies. Weed Res. 22:285294.
341363. San Diego, CA: Academic Press. Vitek, W., and W. Jackson. 1996. Rooted in the land.
Roberts, J. L., and F. R. Olsen. 1942. Interrelation- New Haven, CT: Yale University Press.
ships of legumes and grasses grown in association. Weber, C. R., and D. W. Staniforth. 1957. Competitive
Agron. J. 34: 695701. relationships in variable weed and soybean stands.
Robinson, R. G., and R. S. Dunham. 1954. Compan- Agron. J. 49: 440444.
ion crops for weed control in soybeans. Agron. J. Wiles, L. J., G. G. Wilkerson, and H. J. Gold. 1992.
46:278281. Value of information about weed distribution for
Sakai, K. 1961. Competitive ability in plants: Its improving postemergence control decisions. Crop
inheritance and some related problems. Mecha- Prot. 11:547554.
nisms in biological competition, Soc. for Exp. Biol. Zimdahl, R. L. 1999. Fundamentals of weed science.
Symp. XV. New York: Academic Press. 15:245263. 2d ed. San Diego, CA: Academic Press.
Salisbury, E. J. 1942. The reproductive capacity of
plants. London: G. Bell and Sons.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

Influence of Competition
on the Plant

Interference, the term favored by Harper (1961) to DENSITY

describe interactions among neighboring plants,
Harper and Gajic (1961), studying the response of
strongly affects plant growth, development, and sur-
corn cockle to increasing density, theorized that
vival (Jolliffe 1988). Much of the weed science lit-
plants could respond in two ways: (1) by increased
erature uses the word competition rather than the
mortality, and (2) by increased plasticity in size and
more inclusive term interference to describe the
individual reproductive capacity.
relationships among plants for required environ-
In either way, an individual annual plant can react
mental resources. Competition is used as it was
to increasing density, and thereby its population
defined by Clements et al. (1929) and the weed sci-
becomes self-regulatory. Harper (1964) argued that
ence literature tends to regard it (often correctly) as
the essential properties controlling the ecology of a
the dominant interaction among plants in a commu-
species only can be detected by studying it in com-
nity. Other sources of interference, such as allelopa-
petition, and that its behavior in isolation may be
thy, which has been widely studied by weed
irrelevant to understanding behavior in the commu-
scientists and others, and environmental modifica-
nity. To truly understand the individual in the popu-
tion of competitive interactions, which has been
lation and the population, experimental designs
noted but has not been studied as carefully by weed
must recognize and include the reaction of individu-
scientists, are also important (Jolliffe 1988).
als to the presence of others. Other work has illus-
Sakai (1961) suggested that a plants competitive
trated that density is a highly variable predictor.
ability is a genetic characteristic controlled by poly-
Lutman et al. (1996) asked if early postemergence
genes, whose action is influenced by environmental
assessments of crop and weed vigor could be a more
interactions. Competitive ability can be measured
reliable predictor of weed loss than weed density.
using vegetative growth rate or propagation rate,
They found that predictions based on the relative dry
terms that are usually consistent with each other.
weight of six different crops and wild oats were
However, to be most accurate, plant character varia-
more reliable. It is also interesting that Lutman et al.
tion due to competition must be observed as it is
(1996) found that visual assessments of the potential
affected by intergenotypic competition (Sakai
effects of weeds were quite reliable.
1955). The environment varies in physical attributes
to which plants respond; plants compete for some of
these (water, nutrients, light), but not for others
(e.g., time of planting and time of emergence) Weber and Staniforth (1957) argued that differences
(Sagar 1968). The presence of neighbors of the same in competitive efficiency of crops and weeds are
or different species may alter the environment to well known. This review supports their observation
such a degree that a species that is unable to gain an and provides experimental results to verify it (see
early advantage also may be unable to exploit a chapter 5). However, only a few clues and bits of
competitive advantage later (Sagar 1968). For exam- evidence surface relating to why such things occur.
ple, a high relative growth rate late in the growing While knowing that weeds of a certain species
season becomes valueless if a competitor has con- reduce crop yields a certain percentage is useful, the
sumed the bulk of available soil nutrients. more difficult and interesting question of why this is

20 Chapter 4

true and why the effect varies among crops poses a ability could be predicted from plant traits. Multiple
continuing challenge to research work on plant linear regression showed a strong relationship
interference. between easily observable plant traits and competi-
The competitive ability of four annual weeds and tive ability (r2 = 0.74). Plant biomass explained 63
barley was compared in greenhouse and field stud- percent of the variation in competitive ability, and
ies in the UK (Gustavsson 1986). Wild mustard was plant height, canopy diameter, canopy area, and leaf
consistently more competitive than barley, which shape explained most of the residual variation. Their
was consistently more competitive than field penny- purpose was to encourage use of a predictive tool to
cress, pale smartweed, or scentless chamomile. study competition in natural communities and to
Competitive ability was determined by using rela- encourage plant ecologists to avoid the phenomeno-
tive yields and relative total yield (determined by logical approach to study of competition.
adding the yield of successive cuttings during the The phenomenological approach has also charac-
growing season). Gustavssons (1986) stated pur- terized competition studies done by weed scientists.
pose was to open a discussion on whether or not it For example, Minotti and Sweet (1981) cited sever-
is worthwhile continuing comparisons of the al phenomena as important determinants of the role
growth of species with the method of relative yield of crop interference in limiting losses due to weeds:
and relative total yield. The literature has been silent relative time of emergence and transplanting, the
in response. capacity of crop variety to interfere (its competitive
Studies that describe the outcome of competition ability), allelopathic characteristics, early establish-
do not help us understand if Grime (1979) was right ment of a dense foliar canopy, row spacing and plant
when he proposed that competition was the ten- arrangement, and nutrient and water management.
dency of neighbouring plants to utilize the same Each of these phenomena is important, but their
quantum of light, ion of mineral nutrient, or mole- combination has not yet provided a predictive tool.
cule of water, or volume of space. Nor do they help The literature suffers or is blessed by, depending on
us determine if Tilman (1987) was more correct ones point of view, a large body of special cases but
when he proposed that competition was the utiliza- few general principles. There are literally hundreds
tion of shared resources in short supply by two or of studies that demonstrate that weeds compete with
more species. Competitive ability is then deter- crops (see chapter 5). There is an abundance of lit-
mined by a plants minimum resource requirement, erature that demonstrates that a certain weed densi-
usually designated R*. Descriptive studies also can- ty present in a crop for a certain time will reduce
not tell us if Grace (1990) was correct when he pro- crop yield by a certain amount. General principles
posed that Grime (1979) and Tilman (1987) offered that allow generalizations about weed-crop competi-
complementary not conflicting definitions. Grace tion have not followed from all the preliminary
(1990) suggested that if a habitat is fertile, a species work.
competitive ability is determined by its ability to Clements et at. (1929) described plants competi-
capture the required resources. But if the habitat has tive equipment and provided valuable information
low fertility, competitive ability is determined by a on interplant competition. They cited four points, all
species ability to tolerate low resource availability. centering on the determination of life form:
GENERAL PRINCIPLES 1. Duration or perennationowing its effect on
occupation and height
Gaudet and Keddy (1988) stated that decades of
2. Rate of growthmost effectively expressed
study of interspecific competition in community
by expansion and density of the shoot and
ecology has yielded an overwhelming body of spe-
root systems
cial cases but few general principles. The cause, in
3. Rate and amount of germinationinitial
their view, is the persistent use of the phenomeno-
logical, non-predictive approach. Progress toward
4. Vigor and hardinessfacilitate survival
general principles that allow prediction of competi-
under stress
tive ability from easily observable plant traits
requires a systematic screening of many species Hodgson and Blackman (1956), in a detailed analy-
under standardized conditions. sis of the density response of field bean, concluded
Gaudet and Keddy (1988) used that approach that a profound difference often occurs in the way
with 44 wetland species to determine if competitive plants with determinant and indeterminant growth
Influence of Competition on the Plant 21

respond to density. Species such as field bean, in ly measured and reported weed density. Hume
which the flowering apices do not arise from the major (1985) demonstrated that wheat density varied by as
vegetative apices, mainly respond to density by alter- much as 25 plants m-2 in adjacent quadrats in
ing the number of parts formed. In contrast, common research and farmer field plots.
sunflower and similar species respond more by
Do similar densities have similar effects in all
changes in size of parts. Blackman (1919) studied the
compound interest law and plant growth and stated,
At what stage(s) of development does
what now seems so clear, that in many crop plants the
competition occur?
matter (of plant growth) is of course complicated by
What is the influence of fertility and moisture?
the effect of crowding on the individual plant.
How important is a delay in sowing (or emer-
What are the complicating competitive factors
gence) of the crop in determining the outcome
inherent in the study of plant growth in competition?
of competition?
One reasonable explanation of the sum of the factors
How reproducible are the effects of weeds
encountered by an individual plant was schematical-
from field to field, area to area, and country to
ly outlined by Bleasdale (1960), who proposed that
the competition encountered by an individual plant
depends on the density, distribution, duration, and To this list of questions can be added:
species of its competitors (fig. 4.1). Climatic and
How do different species (or populations) of
edaphic conditions modify the relationships.
weeds compare? (Sagar 1968)
Palmblad (1967, 1968) considered several factors
on the left of Bleasdales (1960) scheme in an inves- There are some clues to answers to these kinds of
tigation of seven weeds. Friesen (1967) posed a questions. The emergence and development of natural
series of questions focusing on the heart of weed- infestations of wild oats was studied in 23 spring bar-
crop competition and enumerating the interactions ley fields in one year and 9 in a second year, in the UK
suggested by Bleasdale (1960). (Peters and Wilson 1983). Wild oats emerged up to the
four-leaf stage of barley and 50 percent emergence
What densities are necessary to reduce yields?
took 22 days in one year and 36 in a second year. The
Do similar densities have similar effects in all
majority of seed was shed by the early emerging
plants in both years. In the first year, 79 percent of the
As affirmed in a brief study by Hume (1985), crop wild oats emerged by barleys two-leaf stage and 59
density is always important as is the more frequent- percent emerged by the two-leaf stage in the second

Figure 4.1.
Schematic dia-
gram depicting
the competition
encountered by
an individual
plant. (Adapted
from Bleasdale
22 Chapter 4

year, and these plants produced 97 and 80 percent of seeding rates by half may not affect crop yield when
all seed shed, respectively (Peters and Wilson 1983). remaining plants more nearly approach their biolog-
Wild oats that emerged before barley produced five ical potential (Harper 1960; Palmblad 1967). Don-
times as much seed per plant as those that emerged ald (1963) said, It is a surprising thought that man,
between the crops two- and three-leaf stage of in growing a successful, healthy field crop creates
growth. Early emerging plants were always the heavi- such intense competition that the individual plants
est, had the most stems per plant, and caused the most are, in a very real sense, subnormal. He obtained
yield loss. maximum levels of dry matter production per pas-
Jennings and Aquino (1968a, b, c) studied the ture plant at lowest densities and observed a
mechanism of competition among rice phenotypes. decreasing trend with increasing density (Donald
Competition was first observed when rice was 53 to 1954). Seeds per inflorescence and the weight per
60 days old, which was 30 to 35 days after trans- seed rose to a peak at intermediate densities, and
planting (1968c). Tall and dwarf cultivars differed then fell.
genetically in ways that affected leaf number, leaf Donalds (1954) reasoning for these results centers
length, leaf angle, and plant height (1968b). Plant on inter- and intraplant competition. With the least-
height had a pronounced effect on yield. Although dense planting, competition was absent during early
short cultivars outyielded tall ones when grown in growth stages when flower primordia originate. As
pure stands, the high-tillering, tall, leafy ones were growth proceeded, interplant competition became pro-
vastly more competitive in mixtures (1968a). Tiller gressively stronger. During flowering and seed forma-
number, number of leaves, leaf length, leaf area tion, the number of inflorescences was so great that
index, height, and dry weight were greater in suc- competition occurred among them. Seed production
cessful competitors, and the differences were efficiency decreased, leading to fewer seeds per inflo-
observable before competition was apparent rescence and reduced seed size, at the widest spacing.
(1968c). Because of their greater height and leaf Therefore, at low densities intraplant competition pre-
area, the better competitors received more light. Tall vails. In extremely dense stands, competition is
genotypes were more competitive under usual already intense when floral primordia originate and
growth conditions and were relatively more compet- both intra- and interplant competition function contin-
itive with added fertility and close spacing. Plant ually. However, in a moderately dense stand, interplant
characteristics (mentioned above, Clements et al. competition operates at the time of floral primordia
1929) that increased size and vegetative vigor dur- initiation reducing the number of floral primordia
ing early growth stages conferred greater competi- formed. This reduced number more nearly matches the
tive ability. plants capacity as interplant competition intensifies,
Donald (1963) stated that plants show extreme while seeds per inflorescence and per unit area achieve
plasticity, responding remarkably in size and form to a maximum. Thus, competition within plants and
environmental conditions. He emphasized that the between plants combine to produce maximum seed
presence of a neighbor constitutes one of the most yield per plant. Donald (1954) suggested that, at low
potent external forces that may limit size and ulti- density, competition within the plant may determine
mate yield. Harper (1964) stated that, The form, the maximum yield of any plant component.
tolerances, and persistence of species may be pro- However, Harper and Gajic (1961) indicated vari-
foundly modified by the proximity of neighbours of ability in seed set was greatest with corn cockle (1
the same or other species. It follows that the charac- to 24 capsules per plant) at low densities and least
teristics of individual species shown by isolated (almost all plants with a single capsule) at high den-
individuals or pure populations may offer no signif- sities. This work suggests that variation may be
icant guidance to their behavior in the presence of greatest at lower densities in contrast to Donalds
others. (1954) findings and other work reported by him
Roots as vital, functional plant parts influence
Characteristics leading to competitive success only competitive relations, although far fewer experi-
can be exposed and demonstrated when species ments have been conducted. Clements (1907) men-
grow together. The concept of plant plasticity in tioned that slight competition occurs between aerial
response to competitive environments was advanced parts of grasses with erect leaves; as a result, com-
by Harper (1960). Because of plasticity, reducing petitive interactions are primarily in the root zone.
Influence of Competition on the Plant 23

Mann and Barnes (1945) thought that with nitrogen obtained and root system lengths decreased 83 to 99
fertilizer in excess of the amount needed by the times.
crop, any yield reduction of barley must be due to Pavlychenko (1937) also grew cereal crops in 6-inch
competition for root space. All possible variables, drill rows with weeds between the rows and compared
with the exception of light, were reportedly more crop root systems 40 days after emergence:
than optimum in the experiment. However, the
weeds, corn spurry and scentless chamomile, are Table 4.1. Comparison of the Root Systems
shorter than barley; hence, light could have been of Barley, Wheat, Wild Oat, and Wild Mustard
Pavlychenko and Harrington (1934, 1935) dis- Ratio of
cussed weeds competitive efficiency in cereal crops Type of competition root system length
and proposed that root system development may be
more important than early germination or the devel- Barleywild oat 7.7 : 1.4
opment of a large assimilation surface; root systems Barleywild oat 6.2 : 2.4
nearest the surface were most effective in competi- Wheatwild mustard 3.3 : 6.5
tion. They found most weeds (20 days after emer- Source: Pavlychenko (1937).
gence) had larger root systems and greater
assimilation surfaces than any of the common cere-
als tested (Pavlychenko and Harrington 1935). In Similar effects were noted in competing aerial
other work, specifically on root development as plant parts.
related to competition, they noted that the capacity Similarly, but in the wetter climate of the UK,
of the root system developed by competitors influ- established perennial ryegrass competed primarily
enced competition between some cereal grains and below ground with invading annual bluegrass and
weeds (Pavlychenko and Harrington 1934). The red fescue (Snaydon and Howe 1986). With root
research was conducted on the western Canadian competition and an absence of nitrogen fertilizer,
plains where moisture is commonly the limiting increasing the density of ryegrass from 2.5 to 40
external factor. Root system capacity immediately plants m-2 reduced the dry weight of invading weeds
after germination and emergence was especially up to 70 times.
important. Further increases in ryegrass density had no
Pavlychenko (1937), in a detailed examination of effect. Red fescue was more affected by ryegrass
the root systems of weeds and crop plants, present- competition than was annual bluegrass. Application
ed a picture of the competitive relations of roots. He of nitrogen fertilizer (400 kg N ha-1) reduced the
traced total root development and carefully mea- effects of root competition.
sured final development. For several days after actu- Black et al. (1969) proposed a biochemical basis
al germination and before emergence, plants for plant competition after examining data from
develop in darkness with no photosynthetic organs. many other studies and placing plants in an efficient
The roots are the main functional exterior organs or nonefficient group based on six criteria. From this
during this period. The size of the plant increases 3 grouping they proposed an hypothesis to explain the
to 400 percent prior to emergence primarily due to competitive success of several crops and weeds. The
root development. Competition, which begins as criteria were:
soon as roots attempt to occupy the same space, may
1. Response to light intensity
occur early in development and affect development
2. Response to increasing temperature
of aboveground parts. Pavlychenko (1937) found
3. Response to atmospheric oxygen
extensive root competition to be the rule. Single
4. Presence of photorespiration
mature plants grown in the center of 10-foot squares
5. Level of photosynthetic carbon dioxide com-
produced total root lengths of:
pensation concentration
Wild oat 3,456,005 inches 6. The pathway of photosynthetic carbon
Wheat 2,802,821 inches dioxide assimilation (C3 versus C4 CO2
Rye 3,114,375 inches fixation)
When the same plants were grown in 6-inch rows After classifying over 50 crops and weeds, they
with 18 to 20 plants per foot, a different ratio was made the logical and not unexpected conclusion that
24 Chapter 4

competition among plants depends on morphology, relative competitive ability of herbicide resistant and
differential response to environmental parameters, susceptible weeds because the phenomenon of resis-
ability to extract nutrients from water and from soil, tance, although known, was not widespread in the
and other factors. However, they proposed that com- late 70s, and the spread of herbicide resistance was
petitive ability also depends on, and partially can be regarded by many as a minor problem. Now it is
explained by, the net capacity to assimilate carbon very important, and the continuing spread of resis-
dioxide and use the photosynthate, an ability intimate- tance is regularly recorded (Heap 2003). In March
ly related to the six criteria. Plants that fix carbon 2003, there were 275 resistant weed biotypes and
dioxide at high rates probably secure an initial com- 165 resistant species (98 dicots and 67 monocots).
petitive advantage and are high-yielding crops or vig- Resistance was present in 59 countries (Heap 2003).
orous weeds. A second paper (Chen et al. 1970) Weaver and Warwick (1982) reported that suscep-
affirmed the validity of the original hypothesis. tible populations of redroot pigweed and Powell
Dakheel et al. (1994) used growth chambers to amaranth had greater competitive ability with
study the effects of three temperature regimes and respect to total biomass and seed production than
two moisture levels on growth, interference, and pho- resistant populations. Their work suggests that resis-
tosynthetic response of downy brome and medusa- tant populations of both species will persist primar-
head. The optimum temperature for growth of both ily in areas where susceptible populations are
species was 24/11C as opposed to 16/5 or 32/16. present at very low densities or have been eliminat-
Moisture limitation reduced yield of both species at ed by regular herbicide application. The relative
all three temperature regimes. Early resource alloca- competitive ability of resistant and susceptible
tion and high growth rates allowed downy brome to plants depends, partially, on the selective pressure
outperform medusahead at high nutrient levels exerted by continued use of the herbicide and on
(Dakheel et al. 1993). These advantages were environmental factors.
reduced when nutrient levels were low; then the Finally, although the thought is now more than 40
species were more nearly equally competitive. The years old, Donald (1963) wrote an excellent and still
two species commonly coexist on rangeland, and valid appraisal of our understanding of competition
these studies showed that in mixtures, with limited or among plants.
unlimited moisture, downy brome had a higher yield
than medusahead in all three temperature regimes. It is a salutary thought that we do not knownor have
Growth chamber studies by Nord et al. (1999) we even given the matter much considerationwhat
determines the density of population of cereal plants
demonstrated that cool spring temperatures give giving maximum yield.Yet until we know this, and espe-
wheat a competitive advantage over kochia and cially until we understand the interaction of density with
Russian thistle. Weaver et al. (1988) proposed that such factors as water and nitrogen, then the develop-
difference in relative times of emergence of crops ment of suitable varieties of plants must depend in the
futureas in the paston empirical plant breeding.
and weeds can be used to suggest optimum planting
We can claim great advances in genetics, and great
times and to estimate potential crop yield losses from advances in producing plants with drought escape or
weed interference. The relative times of emergence disease resistance, fatter pods or finer flowers. And the
of tomato and four weed species at five temperatures breeder can point, too, to varieties which, quite apart
and five soil moisture levels were studied in a growth from these specific virtues, are able under the keen
intraplant competition of a commercial crop, to yield
chamber. In general, total emergence decreased as
more grain, more leaf, more dry matter. Why? The
soil moisture decreased but the species differed in the breeder has no idea. Indeed, the answer to such a
optimum temperature for emergence. The time to 50 question will often be that it yields more because it has
percent emergence decreased with increasing tem- more ears, or more florets or more fertility or less abor-
perature and slightly increased with decreasing soil tion, which of course, is little more than a paraphrase of
the statement that it yields more. Actually, what hap-
moisture. pened was that the breeder selected it because it yield-
The role of water stress is illustrated in studies by ed more, not that it yielded more because it was
Patterson (1986) who showed that growth reduction consciously bred to do so. Why does a modern wheat
due to water stress was greater for soybean than for variety, whether in Greece or New Zealand, yield more
sicklepod. In competition, sicklepod reduced soy- than a variety of like maturity and disease resistance of
fifty years ago? Because it either (a) fixes more carbon
bean growth more when both species were water or (b) has a greater proportion of the carbon in the
stressed than it did when both had adequate water. grain. Why? No one knows. Perhaps it has a different
The first edition of this book did not discuss the root system, better leaf arrangement and light utiliza-
Influence of Competition on the Plant 25

tion, more glume surface, or one of many factors affect- competitive ability of annual weeds. In Proc. Euro-
ing growth and photosynthesis. And, in particular, it has pean Weed Res. Symposium Economic Weed Con-
these desired characteristics when growing under the
trol. European Weed Res. Soc., 105112.
acute stress conditions of a commercial crop.1
Harper, J. L. 1960. Factors controlling plant numbers.
In The biology of weeds, ed. J. L. Harper, 119132.
NOTE Oxford, England: Blackwell Science Pub.
. 1961. Approaches to the study of plant com-
1. Reprinted with permission of Elsevier.
petition. Symp. Soc. For Expt. Biol. 15:139.
. 1964. The individual in the population.
LITERATURE CITED British Ecological Soc. Jubilee Symp., ed. A. Mac-
Black, C. C., T. M. Chen, and R. H. Brown. 1969. fayden and P. J. Newbould, 149158.
Biochemical basis for plant competition. Weed Sci. Harper, J. L., and D. Gajic. 1961. Experimental stud-
17:338344. ies of the mortality and plasticity of a weed. Weed
Blackman, V. H. 1919. The compound interest law Res. 1:91104.
and plant growth. Ann. Bot. (London) 33:353. Heap, I. 2003. International survey of herbicide resis-
Bleasdale, J. K. A. 1960. Studies on plant competi- tant weeds. Accessed
tion. In The biology of weeds, ed. J. L. Harper, March 24, 2003.
133142. Oxford, England: Blackwell Science Pub. Hodgson, G. L., and G. E. Blackman. 1956. An
Chen, T. M., R. H. Brown, and C. C. Black. 1970. analysis of the influence of plant density on the
CO2 compensation concentration, rate of photosyn- growth of Vicia faba. I. The influence of density
thesis and carbonic anhydrase activity of plants. on the pattern of development. J. Exp. Bot.
Weed Sci. 18:399403. 7:147165.
Clements, F. E. 1907. Plant physiology and ecology. Hume, L. 1985. Crop losses in wheat (Triticum aes-
New York: H. Holt and Co. tivum) as determined using weeded and non-
Clements, F. L., J. E. Weaver, and H. C. Hanson. weeded quadrats. Weed Sci. 33:734740.
1929. Plant competition: An analysis of community Jennings, P. R., and R. C. Aquino. 1968a. Studies on
function. Pub. No. 398. Washington, D C: Carnegie competition in rice. I. Competition in mixtures of
Inst. varieties. Evolution 22:119124.
Dakheel, A. J., S. R. Radosevich, and M. G. Barbour. . 1968b. Studies on competition in rice. II.
1993. Effect of nitrogen and phosphorus on growth Competition in segregating populations. Evolution
and interference between Bromus tectorum and Tae- 22:332336.
niatherum asperum. Weed Res. 33:415422. . 1968c. Studies on competition in rice. III.
. 1994. Effects of temperature and moisture on The mechanisms of competition among pheno-
growth, interference and photosynthesis of Bromus types. Evolution 22:529542.
tectorum and Taeniatherum asperum. Weed Res. Jolliffe, P. A. 1988. Evaluating the effects of competi-
34:1122. tive interference on plant performance. J. Theor.
Donald, C. M. 1954. Competition among pasture Biol. 130:447459.
plants. II. The influence of density on flowering Lutman, P. J. W., R. Risiott, and H. P. Osterman.
and seed production in annual pasture plants. Aust. 1996. Investigations into alternative methods to pre-
J Agric. Res. 5:585597. dict the competitive effects of weeds on crop yield.
. 1963. Competition among crop and pasture Weed Sci. 44:290297.
plants. Advances in Agron. 15:1118. Mann, H. H., and T. W. Barnes. 1945. The competi-
Friesen, G. 1967. Weed-crop ecologya science in tion between barley and certain weeds under con-
itself. Sixth Int. Cong. Of Plant Prot. Vienna, Austria. trolled conditions. I. Competition with Spergula
Gaudet, C. L., and P. A. Keddy. 1988. A comparative arvensis Linn. and Matricaria inodora Linn. Ann.
approach to predicting competitive ability from Appl. Biol. 32:1522.
plant traits. Nature 334:242243. Minotti, P. L., and R. D. Sweet. 1981. Role of crop com-
Grace, J. B. 1990. On the relationship between plant petition in limiting losses from weeds. In Handbook of
traits and competitive ability. In Perspectives on pest management in agriculture, vol. 2, ed. D.
plant competition, ed. J. B. Grace and D. Tilman, Pimentel, 351367. Boca Raton, FL: CRC Press, Inc.
5165. San Diego, CA: Academic Press, Inc. Nord, C. A., C. G. Messersmith, and J. D. Nalewaja.
Grime, J. P. 1979. Plant strategies and vegetation 1999. Growth of Kochia scoparia, Salsola iberica
processes. Chichester, UK: J. Wiley and Sons. and Triticum aestivum varies with temperature.
Gustavsson, A. D. 1986. Relative growth rate and Weed Sci. 47:435439.
26 Chapter 4

Palmblad, I. G. 1967. Experimental studies in interfer- Sakai, K. 1955. Competition in plants and its relation
ence in weedy plant species. Diss. Abstr., Sect. B to selection. Cold Spring Harbor Symp. Quant.
27(9):3001-B. Biol. 20:137157.
. 1968. Competition in experimental popula- . 1961. Competitive ability in plants: Its inheri-
tions of weeds with emphasis on the regulation of tance and some related problems. Mechanisms in
population size. Ecology 49:2634. Biological Competition. Soc. for Expt. Biol. Symp.
Patterson, D. T. 1986. Effects of moisture stress on XV. New York: Academic Press, 15:245263.
growth and competitiveness of soybeans and sickle- Snaydon, R. W., and C. D. Howe. 1986. Root and shoot
pod. Abst. Weed Sci. Soc. Am. No. 163. competition between established ryegrass and invad-
Pavlychenko, T. K. 1937. Quantitative study of the ing grass seedlings. J. Appl. Ecol. 23:667674.
entire root system of weed and crop plants under Tilman, D. 1987. Plant strategies and the dynamics of
field conditions. Ecology 18:6279. structure of plant communities. Monographs in
Pavlychenko, T. K., and J. B. Harrington. 1934. Com- Population Biology 20. Princeton, NJ: Princeton
petitive efficiency of weeds and cereal crops. Cana- University Press.
dian J. Res. 10:7794. Weaver, S. E., and S. I. Warwick. 1982. Competitive
. 1935. Root development of weeds and crops relationships between atrazine resistant and suscep-
in competition under dry farming. Scientific Agric. tible populations of Amaranthus retroflexus and A.
16:151160. powellii. from Southern Ontario. New Phytol.
Peters, N. C. B., and B. J. Wilson. 1983. Some studies 92:131139.
on the competition between Avena fatua L. and Weaver, S. E., C. S. Tan, and P. Brain. 1988. Effect of
spring barley. II. Variation of A. fatua emergence temperature and soil moisture on time of emer-
and development and its influence on crop yield. gence of tomatoes and four weed species. Can. J.
Weed Res. 23:305312. Plant Sci. 68:877886.
Sagar, G. R. 1968. Factors affecting the outcome of Weber, C. R., and D. W. Staniforth. 1957. Competitive
competition between crops and weeds. Proc. 9th relationships in variable weed and soybean stands.
Brit. Weed Cont. Conf. pp. 11571162. Agron. J. 49:440444.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

The Effect of Weed Density

The first edition of this book set forth the proposi- A curvilinear relationship was reported by
tion that even a cursory review of a portion of the Roberts and Bond (1975), who described the effect
weed-crop competition literature would lead to the of naturally occurring annual weeds at densities of
conclusion that increasing weed density decreases 65 to 315 plants m-2 on yield of lettuce. Very low
crop yield (Zimdahl 1980). This edition supports densities were not included. Roberts and Bonds
that hypothesis. The weed science literature focuses, (1975) study clearly shows that marketable lettuce
as expected, on the effects of weeds on crops. The yield sinks to zero at less than maximum weed den-
opposite effect is recognized in this chapter, but is sity. It is now clear that the assumptions of the sig-
not a major emphasis of the research reviewed. moidal and curvilinear relationships are incorrect
Mohler (2001) noted that the density, arrangement, and that neither is capable of accurately describing
cultivar, and planting date of the crop that maximize the relationship between weed density and crop
the rate at which the crop occupies space early in the yield (Cousens et al. 1984; Cousens et al. 1985).
growing season usually minimize competitive pres- The relationship is hyperbolic, which means,
sure of weeds. Thus, the effect of crop and weed among other things, that it is best described by a
density is important but the latter is emphasized in hyperbolic curve that is usually referred to as a rec-
the work reviewed for this chapter. tangular hyperbola. It also means that, even though
The first edition did not find many studies of the effect may not be measurable with precision,
weed-crop competition that employed mathematical there is, in theory and actually, an effect of a few
analysis. Weed competition studies had been con- weeds (a low density) on crop yield and growth.
ducted in a wide range of crops by agronomists, hor- Finally, the hyperbolic relationship means that the
ticulturists, and weed scientists who had not made weeds effect cannot exceed total (100 percent)
significant use of mathematical models or mathe- crop loss.
matical description. Clearly, this is no longer true Cousens et al. (1985) point out that the theory of
(see chapter 10). the sigmoidial relationship (Zimdahl 1980) that no
Research has shown that, in most cases, the weed competition threshold exists was based on a faulty
densitycrop yield relationship is not linear. A few statistical assumption and not on sound biological
weeds usually do not affect yield in a way that can principles or on economic rationality. Chapter 10
be detected easily; also, the maximum effect, total includes further discussion of this matter.
crop loss, obviously cannot be exceeded and usual- For all crops reported below, readers are referred
ly occurs at less than the maximum possible weed to the first edition of this book for studies conduct-
density. Based on two assumptions, I proposed in ed prior to 1979. Because of the number of weed-
1980 that weed competition could be represented by crop competition studies in this chapter, they are
a schematic sigmoidal relationship. The assump- arranged alphabetically by crop.
tions were: (1) A few (say five in a hectare) weeds
might affect crop growth and final yield but the Literature Cited
effect could not be measured with any precision, and Cousens, R., B. J. Wilson, and G. W. Cussans. 1985. To
(2) there is a high density of weeds beyond which no spray or not to spray: The theory behind the practice.
further crop yield loss can be measured. British Crop Prot. Conf.Weeds, pp. 671678.

28 Chapter 5

Cousens, R., N. C. B. Peters, and C. J. Marshall. losses, and early harvest minimized but did not
1984. Models of yield lossweed density relation- eliminate losses due to cheat.
ships. In Proc. Seventh Int. Symp. on Weed Biology, Growth-chamber experiments and response sur-
Ecology and Systematics, pp. 367374. face analysis with a linearized hyperbolic equation
Mohler, C. L. 2001. Enhancing the competitive ability of suggested the effects of dandelion on alfalfa were
crops. In Ecological management of agricultural more related to total species density than to the pro-
weeds, ed. M. Liebman, C. L. Mohler, and C. P. Staver, portion of each species in a mixture (Rioux and
269321. New York: Cambridge University Press. Lgre 1992). Duration of competition and alfalfa
Roberts, H. A., and W. Bond. 1975. Combined treat- density (lucerne) accounted for the variation in
ments of propachlor and trifluralin for weed control
shoot biomass of alfalfa grown with dandelion. Dan-
in cabbage. Weed Res. 15: 195198.
delion density explained most of the variation in
Zimdahl, R. L. 1980. Weed-crop competition: A
alfalfa-root biomass.
review. Corvallis, OR: Int. Plant Prot. Center, Ore-
gon State University.
Alfalfa dry matter yield was more influenced by
intra- than by interspecific competition in studies
with bladder campion (Wall and Morrison 1990). In
contrast, the weed was more influenced by interspe-
Alfalfa in weed-free, 55-cm rows produced 820 kg cific competition from alfalfa. Replacement dia-
ha-1 of seed in the year of planting. If a mixed annu- grams and aggressivity indices demonstrated that
al population of 40 weeds in a m of row was present, alfalfa was the dominant and bladder campion the
seed yield was reduced to 45 kg ha-1 (Dawson and subordinate species. Over time the authors expected
Rincker 1982). Competition from 55 weeds m-1 (a alfalfa would become the dominant species. Thus, if
dense stand) of broadleaved weeds reduced yield to bladder campion is present, planting alfalfa is an
80 kg ha-1, whereas a light stand of 4 broadleaved effective weed management technique.
weeds m-1 reduced yield from 820 to 310 kg ha-1.
Literature Cited
Barnyardgrass with 75 culms m-1 of row (a heavy
stand) reduced yield to 160 kg ha-1 (Dawson and Dawson, J. H., and C. M. Rincker. 1982. Weeds in
Rincker 1982). One must conclude that seedling new seedings of alfalfa (Medicago sativa) for seed
alfalfa is not a vigorous competitor. If wheat that production: Competition and control. Weed Sci.
emerged with alfalfa seeded in late August grew for 30:2025.
20 days, alfalfa yield was not affected. When wheat Fischer, A. J., J. H. Dawson, and A. P. Appleby. 1988.
grew more than 30 days, yield decreased and uncon- Interference of annual weeds in seedling alfalfa
trolled wheat reduced first-cutting alfalfa yield more (Medicago sativa). Weed Sci. 36:583588.
than 80 percent (Ott et al. 1990). Ott, P. M., J. H. Dawson, and A. P. Appleby. 1989.
Volunteer wheat (Triticum aestivum) in newly seed-
Similar to all crops, alfalfa yield is depressed
ed alfalfa (Medicago sativa). Weed Technol.
most by annual weeds that emerge with alfalfa and
remain uncontrolled until harvest (Fischer et al.
Pike, D. R., and J. R. Stritzke. 1984. Alfalfa (Medica-
1988). Weeds did not affect alfalfa yield if they were
go sativa)-cheat (Bromus secalinus) competition.
removed before 36 days after alfalfa emergence, a Weed Sci. 32:751756.
figure in close agreement with the work of Ott et al. Rioux, R., and A. Lgre. 1992. Effet de la densit et
(1990). Yield decreased thereafter as the duration of de la proportion des plantes lors de lenvahissement
competition lengthened. Interference was most de la luzerne par Taraxacum officinale Weber. Weed
damaging in early spring when winter annual weed Res. 32:213220.
growth was rapid. Weeds that emerged 65 or more Wall, D. A., and I. N. Morrison. 1990. Competition
days after alfalfa emergence did not affect alfalfa between Silene vulgaris (Moench) Garcke and
yield but often their presence at harvest reduced hay alfalfa (Medicago sativa L.). Weed Res.
quality (Fischer et al. 1988). 30:145152.
In a study with four alfalfa seedling rates in the
fall (4.5, 9, 13.5, and 22 kg ha-1 ) and harvest at three
times (early bud, early bloom, or late bloom), cheat Several papers report studies with both wheat and
(an annual grass) significantly reduced alfalfa pro- barley. In most of these, wheat is the dominant crop,
duction and forage quality (Pike and Stritzke 1984). and these studies are included in the section on
Increasing alfalfa seeding rate only partially offset wheat in this chapter.
The Effect of Weed Density 29

Field experiments conducted in Canada with bar- growth later than barley or wild oats (Cussans and
ley in competition with wild oats showed barley cul- Wilson 1975). Peters and Wilson (1983) studied
tivars differed in their competitive ability. The emergence and development of wild oats in 23
specific cultivars are important to growers and sci- spring barley fields in one year and 9 in a second
entists in the areas where the work was done, but year in the UK. Emergence continued up to the
here the general point of difference is emphasized. crops 4-leaf stage. In one year, the mean number of
Barleys competitive ability (ODonovan et al. days from planting to 50 percent emergence was 22,
2000) always declined when spring emergence was and it was 36 in the second year of the study. The
delayed and always increased when seeding rate majority of seeds shed to create future infestations
increased. Dhima et al. (2000) found, in Greece, that came from early emerging wild oats. In one year, 79
cultivars also differed in competitive ability against percent of the wild oats had emerged by the crops
sterile oat and littleseed canarygrass. Sterile oat at a 2-leaf stage and in the second year the value was 59
density of 120 m-2 was more competitive than 400 percent. These plants contributed 97 and 89 percent
littleseed canary grass m-2. Yield reduction among of the seed shed. Peters and Wilson (1983) were able
four barley cultivars varied from 8 to 67 percent for to establish that wild oats emerging at an early stage
sterile oat, whereas littleseed canarygrass reduced of barley caused a greater yield loss than the same
yield only 1 to 55 percent at the same densities density of later-emerging wild oats. Wild oats that
(Dhima et al. 2000). Torner et al. (1991) reported emerged before the crop produced five times as
that Spanish barley cultivars also differed in their many seeds per plant as those that emerged between
competitive ability with winter wild oat, even when the crops 2- and 3-leaf stage. Early emerging wild
the density of both species was taken into account. oats always caused the greatest yield reduction but
As the density of winter wild oat increased, barley the pattern of emergence varied between years.
yield declined exponentially. Yield declined 10 per- Most wild oats had emerged by the crops 2-leaf
cent with winter wild oat densities of 20 to 80 pani- stage and these produced more than 89 percent of all
cles m-2 and yield losses were 50 percent when seed shed.
panicle density was greater than 300 m-2. In contrast Wilson and Peters (1982) in 51 experiments car-
to the work of Dhima et al. (2000), Torner et al. ried out over 2 years, showed that wild oat infesta-
(1991) found that, in general, barley yield was not tions of spring barley ranging from 8 to 662
affected by barley seeding rate, but the lowest seed- seedlings m-2 resulted in yield reductions of 0 to 72
ing rate (100 kg ha-1) resulted in the highest yield percent. There was a poor relationship between bar-
loss. Low barley seeding rates allowed greater ley yield loss and spring wild oat population. Yield
weed-seed production and assured an infestation in reductions of 0.5 T ha-1 were found from wild oat
future years. Growing season climate was an impor- densities less than 50 seedlings m-2, however yield
tant determinant of competition between barley and losses were up to 1.5 T ha-1 when wild oat density
winter wild oat. Ismail and Hassan (1988) working was greater than 200 plants m-2. Between 50 and 200
in Qatar showed that removing the natural weed seedlings m-2, effects were variable. Yield reductions
population 15, 30, 45, or 60 days after planting pro- were poorly correlated with the number of
duced barley yields similar to a weed-free check. No seedlings, wild oat panicles, or wild oat seed pro-
critical period or threshold density was observed. duced, but there was a good correlation between
Given the number of studies on barley (19 includ- barley yield loss and the dry weight of wild oats pre-
ed), it is clear that wild oats have been regarded as sent at barley harvest. Wilson and Peters (1982) did
the most important weed (12 studies of one of three not find any system of yield loss prediction that
species). In three experiments with barley planted at could be used with confidence.
two rates (90 and 180 kg ha1) and two row widths Evans et al. (1991) used addition series field
(10 and 20 cm), wild oats were more competitive experiments to determine the relative aggressive-
than quackgrass at equivalent shoot populations ness of spring barley and wild oat. Barley was
(Cussans and Wilson 1975). Barley had a greater more aggressive than wild oat, and barley biomass
influence on competition than plant arrangement was more affected by intraspecific competition
(density). The difference between the two weeds whereas wild oat biomass was more affected by
was attributed to their patterns of growth. Wild oats interspecific competition. Increasing wild oat den-
grew much like barley, whereas quackgrass grew sity had a negative, asymptotic-like competition
slowly at first but was able to continue vegetative effect on barley grain yield at all barley densities.
30 Chapter 5

Dunan and Zimdahl (1991) used a replacement emerged at the 0- to 0.5-, 0.5- to 2.5-, or the 2.5- to
series experiment in the field and growth analysis 4-leaf stage of barley achieved densities of 54, 46,
and confirmed the results of Evans et al. (1991) by and 15 m-2. If these densities competed all season,
showing that barley was always the stronger com- seed production was 82, 17, and 1 percent of all seed
petitor. Barleys interspecific competition with shed, respectively, and the yield loss was 16 percent
wild oat was 7.3 times greater than its interspecif- at the highest density but there was no yield loss
ic competition on a dry weight basis. When leaf from the lower densities. Consistent with other stud-
area (see Chapter 9 on methods) was used as the ies reported above, when wild oats were removed up
yield variable, barleys intraspecific competition to barleys 2.5-leaf stage, later emerging wild oats
was only 2.4 times greater than interspecific com- did not compensate by making extra growth. In one
petition owing to barleys higher leaf area. Barley study, barley and wild oat density were 416 and 414
had a greater leaf area, root and shoot biomass, m-2 and in another study in which densities were 295
absolute growth rate, and shoot-root ratio than wild and 294 m-2, grain yield losses were significant only
oat. But wild oat always had a higher leaf area if wild oats remained until barley had 2.5 to 4.5 or
ratio. The relative growth rates (RGR) and net 4.5 to 6.5 leaves, respectively. If a top dressing of
assimilation rates (NAR) did not differ. nitrogen was added when the crop had 3 to 4 leaves,
Morishita and Thill (1988a) found the critical no yield loss occurred unless wild oats remained to
duration of wild oat interference began at about bar- barleys 6-leaf stage (Peters 1984). In a second
leys two-node stage and continued to maturity study, Peters (1985) compared the effects of heavy
when final densities were 160 and 170 plants m-2, (greater than 18 mg) and light (less than 11 mg) wild
respectively. Wild oat reduced barleys biomass, oat seed on barley yield when barley was planted in
number of tillers, tiller heads per unit area, and tiller 25-mm rows. In wild oat grown from heavy seeds
grain yield but not the number or grain yield of bar- sown 75 mm deep with an equal number of barley
leys main stems. Wild oat also did not affect the seed, barley produced 47 percent more panicles, 54
soils matric potential or barleys total plant or the percent more seed, and 56 percent more dry weight
soils nitrogen content. Wild oats presence reduced per plant than plants grown from light seed. When
total water and turgor pressure in barleys boot stage both were sown 25 mm deep, the differences were
and that may affect tiller formation and explain the smaller (21, 28, and 34 percent, respectively). When
results reported. In a separate study, Morishita and barley and wild oat were planted 25 mm deep at
Thill (1988b), using additive culture, showed that equal densities, the dry weight of barley (compared
barley and wild oat tiller head production was to monoculture) was reduced from 10.4 g to 7.7 g
decreased by the presence of either species but plant for heavy seed and was 5.8 g for light seed. When
height was not affected. In mixed culture, wild oats planting depth was 25 mm for barley and 75 mm for
biomass was reduced more at early growth stages wild oat, the weight of a barley plant was reduced
(two to three tillers) than barleys was. They had from 9.5 to 7.2 g by wild oat grown from heavy or
similar total plant nitrogen content throughout the light seed. The reduction in number of grains was
season. Consistent with other studies, Morishita and mainly due to a reduction in the number of fertile
Thill (1988b) showed that barley and wild oats in heads (Peters 1985).
monoculture had similar growth and development. Gonzalez-Ponce (1998) reported on barleys abil-
In mixed culture, barley was more competitive than ity to compete with rigid ryegrass in a replacement
wild oat. In further work, Morishita et al. (1991) series study in the greenhouse. The growth, seed
showed that intraspecific interference affected bar- production, and nitrogen uptake of both species had
ley growth more than interspecific interference from a positive response to nitrogen fertilization. Howev-
wild oat. In fact, interspecific interference from bar- er, nitrogen fertilization did not alter the competitive
ley reduced wild oat growth more than intraspecific relationship between the species. Similar to the rela-
interference among wild oat plants. Nevertheless, tionship between barley and wild oat, barley was the
wild oats are vigorous competitors and emerge more effective competitor, primarily because of its
above the barley canopy near barley anthesis. earlier tillering and greater nitrate absorption.
Peters (1984) studied the growth and competition Conn and Thomas (1987), working in Alaska,
of wild oats that emerged at different times and the showed that common lambsquarters density
time competition began in natural populations of explained 75 percent of the variability in barley yield
spring barley. In one experiment, wild oats that over 2 years. The maximum yield loss attributed to
The Effect of Weed Density 31

common lambsquarters was 23 percent in one year vulgare) and wheat (Triticum aestivum). Weed Sci.
and 36 percent in the second year of the field study. 33:521523.
Elberse and de Kruyf (1979) studied competition Dhima, K. V., I. G. Eleftherohorinos, and I. B. Vasi-
between common lambsquarters and barley with dif- lakoglou. 2000. Interference between Avena sterilis,
ferent dates of emergence in a careful study in the Phalaris minor and five barley cultivars. Weed Res.
Netherlands. When common lambsquarters was 40:549559.
planted 7 days before barley, it did not compete with Dunan, C. M., and R. L. Zimdahl. 1991. Competitive
barley. If the weed was planted 21 or 31 days before ability of wild oats (Avena fatua) and barley
barley, barley could not compete effectively with the (Hordeum vulgare). Weed Sci. 39:558563.
weed. Elberse and de Kruyf (1979) concluded that Elberse, W. Th., and H. N. de Kruyf. 1979. Competi-
light was the most important factor in competition tion between Hordeum vulgare L. and Chenopodi-
um album L. with different dates of emergence of
between these two species. They also concluded that
Chenopodium album. Neth. J. Agric. Sci. 27:1326.
common lambsquarters will not be a detrimental
Evans, R. M., D. C. Thill, L. Tapia, B. Shafii, and J.
weed in barley when it emerges 15 days or less before
M. Lish. 1991. Wild oat (Avena fatua) and spring
barley. The authors properly caution that this work
barley (Hordeum vulgare) affect on spring barley
was done in a controlled climate chamber with opti- grain yield. Weed Technol. 5:3339.
mum conditions for barley growth. The conclusions Gonzalez-Ponce, R. 1998. Competition between barley
therefore may not have application to the field where and Lolium rigidum for nitrate. Weed Res. 38:453460.
conditions for barley growth in the spring are not as Ismail, A. M. A., and A. H. A. Hassan. 1988. Effects
favorable and growing conditions usually favor com- of herbicide timing of removal on interference
mon lambsquarters over barley. between barley and weeds. Weed Res. 28:323330.
OSullivan et al.(1982) developed an index of Morishita, D. W., and D. C. Thill. 1988a. Factors of
competition to measure the interference between wild oat (Avena fatua) interference in spring barley
barley and Canada thistle as a tool to provide an eco- (Hordeum vulgare) growth and yield. Weed Sci.
nomic justification for controlling Canada thistle in 36:3742.
oilseed crops and barley. The model was most reli- . 1988b. Wild oat (Avena fatua) and spring
able when square root transformed weed count data barley (Hordeum vulgare) growth and development
were used with percent yield loss. in monoculture and mixed culture. Weed Sci.
When 30 tartary buckwheat plants m-2 were pre- 36:4348.
sent at barley emergence, barley yield decreased 16 Morishita, D. W., D. C. Thill, and J. E. Hammel.
percent. Yield loss was best represented by a linear 1991. Wild oat (Avena fatua) and spring barley
equation (Hordeum vulgare) interference in a greenhouse
experiment. Weed Sci. 39:149153.

Y = 0.63 + 2.75 x ODonovan. J. T., K. N. Harker, G. W. Clayton, and
L. M. Hall. 2000. Wild oat (Avena fatua) interfer-
ence in barley (Hordeum vulgare) is influenced by
where Y = percent yield loss and x = plant density
barley variety and seeding rate. Weed Technol.
(de St. Remy et al. 1985).
OSullvan, P. A., V. C. Kossatz, G. M. Weiss, and D.
Literature Cited
A. Dew. 1982. An approach to estimating yield loss
Conn, J. S., and D. L. Thomas. 1987. Common barley due to Canada thistle. Can J. Plant Sci.
lambsquarters (Chenopodium album) interference 62:725731.
in spring barley. Weed Technol. 1:312313. Peters, N. C. B. 1985. Competitive effects of Avena
Cussans, G. W., and B. J. Wilson. 1975. Some effects fatua L. Plants derived from seeds of different
of crop row width and seed rate on competition weights. Weed Res. 25:6778.
between spring barley and wild oat (Avena fatua L.) . 1984. Time of onset of competition and
or common couch [Agropyron repens (L.) Beauv.]. effects of various fractions of an Avena fatua L.
In Proc. European Weed Res. Soc. Symp. Status and population in spring barley. Weed Res. 24:305316.
Control of Grassweeds in Europe, pp. 7786. Peters, N. C. B., and B. J. Wilson. 1983. Some studies
De St. Remy, E. A., J. T. ODonovan, A. K. W. Tong, on the competition between Avena fatua L. and
P. A. OSullivan, M. P. Sharma, and D. A. Dew. spring barley. II. Variation of A. fatua emergence
1985. Influence of tartary buckwheat (Fagopyrum and development and its influence on crop yield.
tataricum) density on yield loss of barley (Hordeum Weed Res. 23:305312.
32 Chapter 5

Torner, C., J. L. Gonzalez-Andujar, and C. Fernandez- corn hybrids leaf area index and photosynthetic
Quintanilla. 1991. Wild oat (Avena sterilis L.) com- photon flux receptivity may be useful in developing
petition with winter barley: Plant density effects. integrated weed management systems for corn.
Weed Res. 31:301308. Tillage and row spacing effects were investigated
Wilson, B. J., and N. C. B. Peters. 1982. Some studies by Teasdale (1995). Weed control was poor and
of competition between Avena fatua L. and spring yield was reduced when no herbicides were applied
barley. I. The influence of A. fatua on yield of bar- regardless of row spacing (38 versus 76 cm) or corn
ley. Weed Res. 22:143148. population (standard versus 2x). The corn leaf
canopy in the high population in 38-cm rows
reduced light transmittance 1 week earlier than in
The crop Americans call corn, most of the world the 76-cm rows with a lower plant population. Less
calls maize. The world grows a little less than 140 herbicide (25 percent of recommended rate) provid-
million ha, far less than wheat or rice, but because ed acceptable weed control and grain yields similar
the average yield is greater than 4,200 kg ha-1 , the to the standard treatment with narrow rows and high
total production (590.8 metric tons) is greater than population density but weed control was reduced in
wheat and only slightly less than rice (United 76-cm rows with the low herbicide rate. Murphy et
Nations 2000). Corn is probably the most important al. (1996) showed that increasing corn stand density
crop in the United States in terms of the number of from 7 to 12 plants m-2 or decreasing corn row width
acres and annual value. The United States produces from 75 to 50 cm significantly increased corns leaf
almost one-half of the worlds corn and most is used area index and reduced the photosynthetic photon
for animal feed. flux available to weeds growing below the corn
Begna et al. (2001) found that over two sites and canopy. Narrow rows and high corn density reduced
3 years the decrease in corn biomass production due the biomass of late-emerging weeds. Corn yield
to transplanted and naturally occurring weeds was increased 10 to 15 percent with narrow rows. How-
greater with narrow corn row spacing than it was ever, the gains were not without cost. Intraspecific
from high plant population density. The combina- corn competition in higher density planting reduced
tion of narrow rows and high population densities early corn growth and offset gains in yield from
increased corn canopy light interception 3 to 5 per- reduced weed competition. In comparison to plots
cent. Weed biomass production was five to eight where late-emerging weeds grew without control,
times lower under the corn canopy than when the interrow cultivation did not decrease biomass of
weeds were grown in monoculture. Weed biomass late-emerging weeds and did not increase corn yield.
production was reduced more by early-maturing Only one study (Ford and Mt. Pleasant 1994)
corn hybrids than by late-maturing hybrids with examined the effects of hybrids on competition. The
large leaves. Begna et al. (2001) proposed that six medium-season hybrids differed in leaf angle,
hybrid selection and plant spacing could be used in leaf width, leaf number, leaf area index, height, total
an integrated weed management program. Lindquist dry matter, and grain and stover yield. The lowest
and Mortensen (1998) compared the response of yielding hybrids produced 87 to 91 percent of the
two old and two new corn hybrids. Each hybrid was highest yielding hybrid. In spite of the observed dif-
grown in monoculture and in mixture with velveleaf ferences in hybrids, Ford and Mt. Pleasant (1994)
at 1, 4, 16, or 40 plants m-2. The maximum corn concluded that aboveground plant characteristics
yield loss was 32 percent lower for the two old did not correlate with weed numbers, weed cover, or
hybrids. Velvetleaf capsule production was reduced weed biomass. Their work found a significant inter-
62 percent at low velvetleaf densities with old action between hybrid and weed control for grain
hybrids compared to modern hybrids. In one year, yields in one year, which suggested, but did not
the yield loss of one modern hybrid was 74 percent prove, that some hybrids were more competitive
lower than the other three hybrids at low velvetleaf when weed density was high.
densities, whereas the maximum yield loss of one For the first edition, no reports of velvetleaf com-
old hybrid was 44 percent lower at high density. petition were found, whereas seven are reported here-
Lindquist and Mortensen (1998) found that hybrids in (Zimdahl 1980). Coffman and Frank (1991)
with greater weed tolerance and velvetleaf suppres- reported on a 5-year conservation tillage study in
sive ability had a greater leaf area index, and light Maryland that showed that weed flora had shifted
reception area, which suggests that optimizing a from dominance by giant foxtail to smooth pigweed
The Effect of Weed Density 33

in subsequent years. Annual weeds continue to dom- volume (Bussler et al. 1995) could be used to predict
inate corn studies but they have changed over the individual plant seed production in two separate
years. One suspects the change has not been due to two-species (common cocklebur or velvetleaf versus
the fact that the weeds studied prior to 1979 have dis- corn) interactions.
appeared or that they have been successfully con- Dieleman et al. (1999) found an interaction
trolled. The change in the dominant weed flora and between initial weed seedling density and postemer-
the increasing importance of weeds such as velvetleaf gence herbicide or mechanical weed control for corn
has been due to the response of weeds to the cultural and soybeans in competition with velvetleaf (a range
methods (including control) that have been used. of 0 to 500 seeds m-2) or common sunflower (a range
Such shifts are illustrated by the work of Wilson of 250 to 2,500 seeds m-2). There was a positive lin-
(1993) who evaluated the effect of tillage and herbi- ear relationship between initial weed seedling densi-
cides on weed density and corn grain yield over 4 ty and the density of surviving seedlings. Weed
years in Nebraska. Ridge-tillage favored develop- management outcomes for a range of management
ment of kochia and reduced density of wild proso intensities were dependent on initial seedling densi-
millet and common lambsquarters. Tandem disking ty. As initial density increased, the absolute number
increased longspine sandbur and redroot pigweed of survivors increased but the proportion of sur-
density, whereas moldboard plowing favored com- vivors appeared to remain constant over the density
mon sunflower. Cultivation of the crop reduced ranges included. Their results emphasize the need,
weed density 86 percent, but the remaining weeds in any crop, to know the weed density when assess-
reduced corn yield 40 percent compared to a hand- ing the efficacy of any weed management system.
weeded control. Wilson (1993) advocated recogni- Ayeni et al. (1984a,b,c) studied weed interference
tion of the effects of land preparation on weed in corn and cowpea and a corn-cowpea intercrop in
population and the integration of cultivation and Nigeria. They found that weed interference effects
herbicides in weed management strategies. on crops under no-tillage depended on cropping sea-
The effects of conventional tillage and no-tillage son (year), cropping pattern, and crop species
on the outcome of early weed competition in corn (Ayeni et al. 1984c). Except for the corn-cowpea
was studied in Nigeria by Ayeni et al. (1984a). There intercrop that showed significant yield reduction
were more different weedy species with no-tillage, when exposed to 4 weeks of weed interference early
but total weed weight was only 52 percent of the in the season, all cropping patterns required more
weed weight in conventionally tilled plots 6 weeks than 4 weeks of weed interference before yield
after corn planting. Cropping pattern had no effect reduction could be measured. Weed interference was
on plot weediness. With minimum or no weed inter- more detrimental to yield in monocultures in the
ference, corn yield was better after conventional early season than was true in the intercrop. Late in
than after no-tillage but worse if weeds were the growing season, all cropping patterns were
allowed to grow well into the season. equally sensitive to weed interference. In the first 6
In work in Colorado (VanGessel 1995), weed den- weeks of growth, cropping pattern had no effect on
sities of 0, 33, 50, or 100 percent of the indigenous weed growth. Weeds did not affect crop growth until
population affected corn yield but the weeds distri- 5 to 6 weeks after crop planting (Ayeni et al. 1984b).
bution in the field did not. Each additional weed Total crop dry weight was not affected by cropping
reduced corn yield 8.5 kg ha-1 in one year and 2.3 kg pattern. Three weeks after crop planting, weeds
ha-1 in the second year of the study. Weed density 5 from weedy, cropped plots had taken up two to four
to 8 weeks after planting provided a better estimate times as many nutrients (N, P, K, Ca, and Mg) as
of the eventual effect of weeds on yield than did weed-free crops. Remison (1979), also working in
weed density immediately before corn harvest. Bus- Nigeria, found no significant interaction between
sler et al. (1995) used aboveground plant volume to weeding and nitrogen at any site. After corn had
quantify interference of common cocklebur and vel- been grown for several years, weed competition
vetleaf with corn. They found that the ratio of lowered corn yield as much as 50 percent, but two
aboveground plant volume to the total neighborhood hand-weedings 3 and 7 weeks after planting were as
volume was the independent variable that accounted efficacious as more intensive weeding.
for the most variation in target plant seed produc- In recent years, several studies have been done to
tion. Although the method must be verified in dif- determine the effect of competition on growth and
ferent soil, fertility, and water environments, plant seed production of some weeds. Longspine sandbur
34 Chapter 5

has become a problem in corn in the Great Plains. If Work by Frantik (1994) showed that competition
it emerges in late May, with or soon after corn, from all weeds is not equal. Chenopodium suecicum
seedlings produce 1,120 burs per plant. Seedlings J. Murr. and redroot pigweed were competitively
emerging 4 weeks later produce 84 percent fewer equivalent to 1.1 and 0.26 corn plants, respectively,
burs (Anderson 1997). If longspine sandbur is con- whereas 1 corn plant was equal to 5.5 weed plants as
trolled before 4 weeks of interference occur, there is measured by effects on biomass. Redroot pigweed
no loss of corn grain yield (Anderson 1997). Bur- was a more effective competitor with corn and with
cucumber seed production was studied by Esben- C. suecicum than the reverse. Corn reduced the com-
shade et al. (2001). Burcucumber grown without petitive effectiveness of redroot pigweed against C.
competition from corn produced 716 g dry matter suecicum. Corn yield loss increased as the propor-
and 4,500 seeds per plant. Biomass was greatest for tion of C. suecicum in the mixture increased. C. sue-
plants established in May but seed production was cicum that was seeded before corn and not removed
greatest for those established in mid-June. When until 32 days after corn emergence significantly
burcucumber established in corn, it produced 96 reduced corn yield (Frantik 1994).
percent less dry matter and seed than plants grown Several papers report specific effects of specific
without competition, but seed was still produced. weeds on corn yield reduction. More than 13 species
The importance of velvetleaf in corn is empha- have been studied and the papers are summarized
sized by the work of Scholes et al. (1995) who found below and in table 5.1.
that when velvetleaf was grown in corn at densities Natural stands of hemp dogbane reduced corn
of 0, 1.3, 4, 12, or 24 plants m-2, the weeds leaf area yield not at all or up to 10 percent (Schultz and
index and total plant biomass were correlated with Burnside 1979). Jimsonweed decreased corn yield
velvetleaf density and both were negatively correlat- 14 to 63 percent when corn density was 8.3 plants
ed with corn biomass. The maximum yield loss was m-2 and jimsonweed density was 8.3 or 16.7 plants
37.2 percent with a loss of 4.4 percent per unit of m-2, in Spain (Cavero et al. 1999). Yield reduction,
velvetleaf density. as one would expect, increased as the time between
Cardina et al. (1995) found that corn yield loss crop and weed emergence decreased. Corns leaf
and velvetleaf seed production were higher in a area developed faster and it was able to successfully
warm-wet year than in a dry-cold year. Corn yield compete for light with jimsonweed, although the
loss was generally greater with no-tillage than in weeds competitive effects on corn (reduced crop
conventional tillage and from early rather than late- growth rate, lower grain number per ear, and
maturing velvetleaf. The maximum velvetleaf seed reduced grain weight) could be observed late in the
production ranged from 18,000 seeds m-2 for early growing season. When giant ragweed and corn
emerging weeds in no-till to only 100 seeds m-2 for emerged concurrently, densities of 1.7, 6.9, and 13.8
late-emerging weeds. Corn reduced velvetleafs weeds 10 m-2 gave a predicted loss rate of 13.6 per-
seed rain by 50 percent (Zanin and Sattin 1988). cent for the first weed 10 m-2 in the linear response
Seed rain reached a maximum at 20 to 30 plants range at low densities and a maximum loss of 90
m-2 in corn, and 30 to 35 plants m-2 in monoculture. percent at high weed densities (Harrison et al.
When only 4 to 5 velvetleaf m-2 competed with 2001). Corn yield loss response was linear when
corn, the weed produced eight thousand to ten thou- giant ragweed emerged 4 weeks after corn and it
sand seeds, indicative of the weeds great seed pro- was equivalent to a yield loss rate of 1 percent per
duction potential. Zanin and Sattin (1988) also unit increase in weed density. Giant ragweeds seed
calculated the economic threshold to be between 0.3 production decreased significantly with delayed
and 2.4 weeds m-2 for velvetleaf in corn based on emergence. For common milkweed, corn yield loss
studies that included weed densities from 0 to 80 m-2. ranged from 2 to 10 percent with milkweed densities
Defelice et al. (1988) found that monocultural between 11,000 and 45,200 plants ha-1. (Cramer and
velvetleaf had a greater dry weight, leaf area index, Burnside 1982).
and height than when it grew with corn. Velvetleaf When Palmer amaranth was grown with irrigated
planted 5 weeks after corn was lower in all indices corn at densities of 0, 0.5, 1, 2, 4, or 8 plants m-1 of
than that planted with corn. There was no effect of row and the weed emerged with corn, corn yield
tillage system on velvetleaf plant characteristics. declined 11 to 91 percent as Palmer amaranth densi-
Interference from corn and delayed planting reduced ty increased from 0.5 to 8 plants m-1 (Massinga et al.
velvetleaf population at the end of the season. 2001). However, when the weed emerged later than
The Effect of Weed Density 35

Table 5.1. Effect of Several Weeds on Corn Yield

Weed Species Density Yield reduction Source

Barnyardgrass 100 m2 18% Kropff et al. 1984

200 m2 concurrent emergence 2635% Bosnic et al. 1997
Emergence when corn 6%
had 4 leaves

Common milkweed 11,000 to 2 to 45,200 10% Cramer and Burnside

plants m2 1982

Giant ragweed 1.7, 6.9, or 13.6% Harrison et al. 2001

13.8 plants m2 90%

Giant foxtail 10 m1 1314% Fausey et al. 1997

Green foxtail 0, 29, 56, 2056% Sibuga and Bandeen

or 89 m2 but nonsignificant 1980
129 m2 5.817.6%
Hemp dogbane Natural 010% Schultz 1979

Itchgrass 2, 4, up to 14 wk 125 kg ha-1 for each Strahan et al. 2000

week of presence
Season-long 33%

Jimsonweed 8.3 or 16.7 1463% plants m-2 Cavero et al. 1999

Quackgrass 65 to 390 shoots m2 1216% Young et al. 1984

745 shoots m2 37%

Palmer amaranth 0.5 to 8 m2 1174% Massinga et al. 2002

Redroot pigweed 0.5 m-1 with 5% Knezevic et al. 1994

concurrent planting or 4 m1
with planting at corns 35
leaf stage
Wild proso millet 10 m2 1322% Wilson and Westra 1991

Yellow nutsedge 100 shoots m2 8%/100 shoots Stoller et al. 1979

300 tubers m2 17%
700 tubers m2 41%

corn, yield loss occurred only when the weed and forage quality of weeds harvested with corn
emerged at corns 4- and 6-leaf stages. Palmer ama- both declined with increasing Palmer amaranth den-
ranths seed production per plant decreased with sity (Massinga and Currie 2002). Forage yield
greater density, but seed per unit area increased from declined 1 to 44 percent of the weed-free yield with
140,000 to 514,000 seeds m-1 at 0.5 and 8 plants m-1, Palmer amaranth densities of 0.5 to 8 plants m-1 of
respectively (Massinga et al. 2001). When Palmer row, whereas grain yield declined 11 to 74 percent at
amaranth grew with irrigated corn at densities of 0, the same weed densities. Thus, Palmer amaranth
0.5, 1, 2, 4, or 8 plants m-1 of row, corn grain yield interference in corn may not affect forage quality
36 Chapter 5

much, but corn yields decline significantly (Massin- year and 14 percent in a second year from 10 giant
ga 2002). Redroot pigweed performed similarly at foxtail m-1 of row. Corn dry matter at maturity was
the same densities (Knezevic et al. 1994). When red- reduced by nearly one-quarter from the same densi-
root pigweed and corn were planted concurrently, ties. Ten weeds in a meter of row produced 15,700
0.5 weeds m-1 of row reduced corn yield 5 percent, seeds, and their germination was not affected by
which was the same reduction obtained from 4 plant density (Fausey et al. 1997).
weeds when they were planted at corns 3- to 5-leaf Barnyardgrass has been a common weed in sever-
stage of growth. Redroot pigweed that emerged after al crops for decades. A natural stand of barnyard-
corn had 7 leaves did not decrease corns yield. grass with an average density of 100 weeds m-2
Therefore, Knezevic et al. (1994) concluded that the reduced corn yield to just 18 percent of the weed-
time of the weeds emergence was more important free control in the Netherlands. The yield reduction
than its density. varied a great deal between years primarily due to
Sibuga and Bandeen (1980) studied full-season differences in the relative emergence time of the
interference from common lambsquarters and green crop and the weed and the resultant competition for
foxtail. Nonsignificant yield reductions were light (Kropff et al. 1984). Bosnic and Swanton
obtained from green foxtail densities of 20 and 56 (1997) also reported the importance of relative time
plants m-2. Increased green foxtail densities of 89 of emergence. When barnyardgrass and corn were
and 129 plants m-2 reduced corn yield 5.6 to 17.6 planted at the same time and when barnyardgrass
percent over 2 years. Common lambsquarters densi- was planted at the 1- to 2- or 3- to 4-leaf stage of
ty less than 109 m-2 did not affect corn yield over 2 corn growth, yield losses were quite different. Barn-
years. Densities greater than 172 m-2 reduced yield yardgrass at a density of 200 plants m-2 and planted
12 to 38 percent one year and 6 to 58 percent in a with corn reduced yield 26 to 35 percent but less
second year. The effects of the two weeds were sim- than 6 percent if it emerged after corn had four
ilar. They differed primarily in the ability of com- leaves. Barnyardgrass that emerged up to corns 3-
mon lambsquarters to reduce corn ear and seed size leaf stage produced an average of 34,600 seeds m-2
(Sibuga and Bandeen 1980). whereas barnyardgrass that emerged after corn had
Beckett et al. (1988) compared corn yield reduc- four leaves produced only 1,200 to 2,800 seeds m-2.
tion caused by season-long interference from shat- Mickelson and Harvey (1999) determined the
tercane, common lambsquarters, common effects of density and time of emergence on woolly
cocklebur, and giant foxtail at densities of 0.4 to cupgrass growth and seed production in corn. Com-
13.1 plants or clumps m-1 of row. Corn seed yield pared to woolly cupgrass grown at 3 plants m-2 that
decreased linearly with increasing density of 2 to 3 emerged with corn, total aboveground mature bio-
clumps of shattercane or 5 to 8 clumps of giant fox- mass was reduced 54, 97, and 99 percent when
tail. There was a 22 percent yield loss from 6.6 shat- woolly cupgrass emerged at the V2, V5, or V10
tercane clumps m-1 of row, a 27 percent loss from stages of corn growth in one year and by 70, 87, and
4.7 common cocklebur m-1 of row in one year and a 99 in a second year. Woolly cupgrass aboveground
10 percent loss from 6.6 common cocklebur m-1 in 2 vegetative biomass production per mature plant was
of 3 years. Common lambsquarters reduced yield in linearly related to seed production per plant, which
only one year, and the maximum loss was 12 percent decreased nonlinearly as density decreased and time
from 4.9 m-1 of row. There was an 18 percent loss in of emergence was delayed (Mickelson and Harvey
corn yield from 13.1 giant foxtail clumps m-1 of row 1999). Seed production was 12,700 per plant in one
(Beckett et al. 1988). year and 57,100 in the other. The work suggests that
There is no reason to argue that broadleaved late-emerging woolly cupgrass is not of great impor-
species are more detrimental or more ubiquitous tance to long-term management of the weed.
than annual grass weeds in corn. Because the stud- Wild proso millet is an important weed across the
ies have been done in so many different places in semiarid Great Plains of the United States, where
different years, it is not possible to say what type of corn may be grown in rotation with winter wheat
weed is more detrimental. Several studies have and proso millet (mainly for bird seed). In corn, wild
reported on interference from grass weeds. proso millet seedlings begin to emerge in May in
Fausey et al. (1997) studied giant foxtail interfer- Colorado within 2 weeks of corn emergence. Early
ence in corn at densities of 0, 10, 20, 30, 60, 84, or emerging seeds produced the most seeds (2,800 per
98 m-1. Corn yield was reduced 13 percent in one plant), whereas seedlings that emerged 4 weeks later
The Effect of Weed Density 37

produced 88 percent fewer seeds (Anderson 2000). yellow nutsedge is a key to effective yellow nut-
If the weed was controlled by late June, there was no sedge management (Ghafar and Watson 1983b).
loss of corn grain yield (Anderson 2000).
Itchgrass did not compete at all with corn or was Literature Cited
allowed to compete for 2, 4, 6, 8, 10, 12, or 14 (all Anderson, R. L. 2000. Ecology and interference of
season) weeks. Season long (14-week) interference proso millet (Panicum miliaceum) in semi-arid
reduced corn height 18 percent and yield 33 percent corn. Weed Technol. 14:4550.
over 2 years. Each week of interference reduced . 1997. Longspine sandbur (Cenchrus
corn yield 125 kg ha-1. Itchgrass competition for loongispinus) ecology and interference in irrigated
more than 2 weeks after corn emergence always corn (Zea mays). Weed Technol. 11:667671.
reduced corn yield (Strahan et al. 2000). Wild proso Ayeni, A. O., W. B. Duke, and I. O. Akobundu. 1984a.
millet is a more ubiquitous weed across the U.S. Weed interference in maize, cowpea, and
central Great Plains, and although it is not as large a maize/cowpea intercrop in a subhumid tropical
plant as itchgrass, it is as effective a competitor. Wil- environment. I. Influence of cropping season. Weed
son and Westra (1991) found yield reductions of 13 Res. 24:269280.
to 22 percent from 10 wild proso millet plants m-2. If . 1984b. Weed interference in maize, cowpea,
and maize/cowpea intercrop in a subhumid tropical
weed removal was delayed for 2 weeks after corn
environment. II. Early growth and nutrient content
planting, yield could be reduced as much as 10 per-
of crops and weeds. Weed Res. 24:281290.
cent. If removal was delayed up to 6 weeks after
. 1984c. Weed interference in maize, cowpea,
planting, corn yield was reduced 16 to 28 percent
and maize/cowpea intercrop in a subhumid tropical
(Wilson and Westra 1991). environment. III. Influence of land preparation.
Quackgrass is a vigorous competitor with corn Weed Res. 24:439448.
and reduced yield 12 to 16 percent with 65 to 390 Beckett, T. H., E. W. Stoller, and L. M. Wax. 1988.
shoots m-2. Yield reduction was 37 percent with 745 Interference of four annual weeds in corn (Zea
shoots m-2 (Young et al. 1984). Young et al. (1984) mays). Weed Sci. 36:764769.
demonstrated that when light and nutrients were not Begna, S. H., R. I. Hamilton, L. M. Dwyer, D. W.
limiting, an adequate supply of soil water can elim- Stewart, D. Cloutier, L. Assemat, K. Foroutan Pour,
inate the detrimental effects of quackgrass on corn. and D. L. Smith. 2001. Weed biomass production
When soil water was limiting, irrigation increased response to plant spacing and corn (Zea mays)
the yield of quackgrass-free corn and of quackgrass- hybrids differing in canopy architecture. Weed
infested corn. The presence of quackgrass did not Technol. 15:647653.
affect corns nutrient status (Young et al. 1984). Bosnic, A. C., and C. J. Swanton. 1997. Influence of
Stoller et al. (1979) showed that corn yield was barnyardgrass (Echinochloa crus-galli) time of
reduced 8 percent for every 100 yellow nutsedge emergence on density of corn (Zea mays). Weed
shoots m-2. Early competition was the most detri- Sci. 45:276282.
mental and use of preplant herbicides always Bussler, B. H., B. D. Maxwell, and K. J. Puettmann.
reduced yield loss. Without control, corn yield 1995. Using plant volume to quantify interference
declined 17 percent with 300 yellow nutsedge tubers in corn (Zea mays) neighborhoods. Weed Sci.
m-2 and 41 percent with 1,700 tubers m-2. The per-
Cavero, J., C. Zaragoza, M. L. Suso, and A. Pardo.
sistence of the tubers is emphasized by their finding
1999. Competition between maize and Datura stra-
that at least 2 years of effective control were
monium in an irrigated field under semi-arid condi-
required to reduce tuber numbers to 20 percent of
tions. Weed Res. 39:225240.
the original number and 3 years to gain an addition- Cardina, J., E. Regnier, and D. Sparrow. 1995. Vel-
al 5 percent reduction. Ghafar and Watson (1983a) vetleaf (Abutilon theophrasti) competition and eco-
recommended the use of crop density to manage nomic thresholds in conventional and no-tillage
yellow nutsedge. When corn density increased from corn (Zea mays). Weed Sci. 43:8187.
33,300 to 133,200 plants ha-1, yellow nutsedges Coffman C. B., and J. R. Frank. 1991. Weed-crop
aboveground biomass, tuber numbers, tuber weight, responses to weed management systems in conserva-
and height all declined at the end of the growing sea- tion tillage corn (Zea mays). Weed Technol. 5:7681.
son, primarily due to increased light competition Cramer, G. L., and O. C. Burnside. 1982. Distribution
from the more dense corn stand. Manipulation of the and interference of common milkweed (Asclepias
corn planting date to assure that it emerged prior to syriaca) in Nebraska. Weed Sci. 30:385388.
38 Chapter 5

Defelice, M. S., W. W. Witt, and M. Barrett. 1988. growth and seed production in Zea mays. Weed Sci.
Velvetleaf (Abutilon theophrasti) growth and devel- 47Z:687692.
opment in conventional and no-tillage corn (Zea Murphy, S. D., Y. Yakubu, S. F. Wiese, and C. J.
mays). Weed Sci. 36:609615. Swanton. 1996. Effect of planting patterns and
Dieleman, J. A., D. A. Mortensen, and A. R. Martin. inter-row cultivation on competition between corn
1999. Influence of velvetleaf (Abutilon theophrasti (Zea mays) and late emerging weeds. Weed Sci.
L.) and common sunflower (Helianthus annuus) 44:856870.
density variation on weed management outcomes. Remison, S. V. 1979. Effects of weeding and nitrogen
Weed Sci. 47:8189. treatments on maize yields in Nigeria. Weed Res.
Esbenshade, W. R., W. S. Curran, G. W. Roth, N. L. 19:7174.
Hartwig, and M. D. Orzolek. 2001. Effect of estab- Scholes, C., S. A. Clay, and K. Brix-Davis. 1995. Vel-
vetleaf (Abutilon theophrasti) effect on corn (Zea
lishment date and crop competition on burcucum-
mays) growth and yield in South Dakota. Weed
ber fecundity. Weed Sci. 49Z:524527.
Technol. 9:665668.
Fausey, J. C., J. J. Kells, S. M. Swinton, and K. A.
Schultz, M. E., and O. C. Burnside. 1979. Distribu-
Renner. 1997. Giant foxtail (Setaria faberi) inter-
tion, competition and phenology of hemp dogbane
ference in nonirrigated corn (Zea mays). Weed Sci.
(Apocynum cannabinum) in Nebraska. Weed Sci.
45:256260. 27:565570.
Ford, G. T., and J. Mt. Pleasant. 1994. Competitive Sibuga, K. P., and J. D. Bandeen. 1980. Effects of
abilities of six corn (Zea mays) hybrids with four green foxtail and lambs-quarters interference in
weed control practices. Weed Technol. 8:124128. field corn. Can. J. Plant Sci. 60:14191425.
Frantik, T. 1994. Interference of Chenopodium sueci- Stoller, E. W., L. M. Wax, and F. W. Slife. 1979. Yel-
cum J. Murr. and Amaranthus retroflexus L. in low nutsedge (Cyperus esculentus) competition
maize. Weed Res. 34Z:4554. and control in corn (Zea mays). Weed Sci.
Ghafar, Z., and A. Watson. 1983a. Effect of corn (Zea 27:3237.
mays) population on the growth of yellow nutsedge Strahan, R. E., J. L. Griffin, D. B. Reynolds, and D.
(Cyperus esculentus). Weed Sci. 31:588592. K. Miller. 2000. Interference between Rottboelllia
. 1983b. Effect of corn (Zea mays) seeding cochinchinensis and Zea mays. Weed Sci.
date on the growth of yellow nutsedge (Cyperus 48:205211.
esculentus). Weed Sci. 31:572575. Teasdale, J. R. 1995. Influence of narrow row/high
Harrison, S. K., E. E. Regnier, J. T. Schmoll, and J. E. population corn (Zea mays) on weed control and
Webb. 2001. Competition and fecundity of giant light transmittance. Weed Technol. 9:113118.
ragweed in corn. Weed Sci. 49:224229. United Nations Food and Agriculture Organization.
Knezevic, S. Z., S. F. Wiese, and C. J. Swanton. 1994, 2000. Production yearbook 54: 7475.
Interference of redroot pigweed (Amaranthus VanGessel, M. J., E. E. Schweizer, K. A. Garrett, and
retroflexus) in corn (Zea mays). Weed Sci. 42:568573. P. Westra. 1995. Influence of weed density and dis-
Kropff, M. J., F. J. H. Vossen, C. J. T. Spitters, and W. tribution on corn (Zea mays). Weed Sci.
de Groot. 1984. Competition between a maize crop
Wilson, R. G. 1993. Effect of preplant tillage, post-
and a natural population of Echinochloa crus-galli
plant cultivation, and herbicides on weed density in
(L.). Neth. J. Agric. Sci. 32:324327.
corn (Zea mays). Weed Technol. 7:728734.
Lindquist, J. L., and D. A. Mortensen. 1998. Toler-
Wilson, R. G., and P. Westra. 1991. Wild proso millet
ance and velvetleaf (Abutilon theophrasti) suppres-
(Panicum miliaceum) interference in corn (Zea
sive ability of two old and two modern corn (Zea mays). Weed Sci. 39:217220.
mays) hybrids. Weed Sci. 48:569574. Young, F. L., D. L. Wyse, and R. J. Jones. 1984.
Massinga, R. A., and R. S. Currie. 2002. Impact of Quackgrass (Agropyron repens) interference on
Palmer amaranth (Amaranthus palmeri) on corn corn (Zea mays). Weed Sci. 32:226234.
(Zea mays) grain yield and yield and quality of for- Zanin, G., and M. Sattin. 1988. Threshold level and
age. Weed Technol. 16:532536. seed production of velvetleaf (Abutilon
Massinga, R. A, R. S. Currie, M. Horak, and J. Buyer, theophrasti Medicus) in maize. Weed Res.
Jr. 2001. Interference of Palmer amaranth in corn. 28:347352.
Weed Sci. 49:202208. Zimdahl, R. L. 1980. Weed-crop competition: A
Mickelson, J. A., and R. G. Harvey. 1999. Effects of review. Corvallis, OR, Int. Plant Prot. Center, Ore-
Eriochloa villosa density and time of emergence on gon State Univ. 196 pp.
The Effect of Weed Density 39

COTTONGOSSYPIUM HIRSUTUM L. ranged from 3 to 27 percent. Common cocklebur,

jimsonweed, and common ragweed reduced cotton
Cotton began to replace wool as peoples most yield 28, 15, or 12 percent, respectively. Spurred
important fiber in the nineteenth century. China has anoda and common cocklebur exerted an influence
been the worlds major producer and the United 160 and 136 cm from the cotton row, whereas sick-
States has been a close second. Cotton in Europe, lepod had an influence of only 46 cm. The primary,
Asia, and Africa probably originated in what is now but not startling, conclusion one can draw from this
Pakistan (Hobhouse 1985, p. 142). Cotton may have work is that the influence of a weed or an aggregate
been endemic in the New World but how it got to the of weeds is highly variable but all have some effect.
United States remains a mystery. One bale of U. S. No general principles or generalizations can be
cotton was exported to Liverpool in 1784. It was not derived except what I have earlier (see preface)
accepted and rotted on the pier because it had not called the central hypothesis of Weed Science:
entered on a British ship. From that bad beginning, Weeds compete with crops and reduce crop yield
the cotton trade grew to 4 million bales, and the his- and quality.
tory of the growth of that part of U.S. agriculture Many studies in support of this hypothesis have
includes the whole history of Southern slavery and been reported in the last several years. Studies on 30
the Civil War (Hobhouse 1985, p. 142). different weeds are summarized below but no addi-
The interference of more than 30 different weed tional general principles can be derived from these
species has been studied in cotton. A general survey papers. Keeley and Thullen spent a great deal of
in California (Kempen 1984) reported that several time and effort in a quest to analyze the competi-
annual and perennial weeds reduced cotton lint pro- tiveness of all weeds important in cotton in Califor-
duction by 0.5 bale A-1 or more. More than 50 per- nia. Their work on several weed species will be cited
cent of Kern County, CA, cotton fields were weed frequently below in studies with similar design and
free at harvest. In weedy fields, the dominant objectives.
species varied between clay and sandy soils with
BarnyardgrassEchinochloa crus-galli
some causing severe, some moderate, and some only
light losses. Barnyardgrass that competed for 6, 9, 12 or 25
Planting date had little influence on weed inter- weeks after cotton emergence reduced cotton yield
ference in cotton in Arkansas. Losses increased 30 21, 59, 90, or 97 percent. A weed-free period of 9
to 50 percent when weed density increased from 1.7 weeks after emergence was required to prevent cot-
to 6.7 m-1 of row (Klingaman and Oliver 1994). ton yield reduction. If cotton was kept weed free for
Miller et al. (1983) found no yield advantage for 51- 3 or 6 weeks after emergence, it yielded 13 and 87
or 102-cm rows in California. Barnyardgrass densi- percent as much as cotton that was weed free for the
ty increased after cotton layby regardless of row whole season (Keeley and Thullen 1991a). The
width, so that at harvest the number of weeds was importance of regular control is illustrated by the
equal for the two row widths. Similarly, cotton fact that barnyardgrass produced 80 to 90 percent
yields were not affected consistently by interrow fewer seeds in plots kept weed free for the first 6
cultivation in a four-state, 3-year study (Colvin et al. weeks after emergence. Weeds that emerged after 9
1992). When all frequencies were considered, culti- to 12 weeks of weed control did not grow more than
vation initiated 1 or 2 weeks after cotton emergence 10 cm tall due to shading by cotton and did not pro-
and continued for two, four, or six times increased duce seed. Keeley and Thullen (1991a) proposed
yield at only three of nine locations over all years. that barnyardgrass is as competitive as weeds known
For two locations, cotton yields were increased by to be particularly aggressive such as johnsongrass.
only two cultivations.
BermudagrassCynodon dactylon
Byrd and Coble (1991b) compared the amount of
cotton yield loss caused by eight annual weeds, each In short-term competition experiments done in the
at a density of one weed in each 3 m of row. Yields greenhouse, growth of cotton planted 3 weeks after
were reduced between 1 and 7 percent. Sicklepod bermudagrass was severely reduced to about 15
did not cause any yield loss, while redroot pigweed, percent of the control 10 weeks after planting. Cot-
common cocklebur, and common ragweed ton growth was not affected when it was planted
decreased yield 7, 6, and 5 percent, respectively. In before the weeds (Horowitz 1973). The effect was
a second year, yield losses from individual weeds attributed to the weeds rapid growth and possible
40 Chapter 5

allelopathic effects. Similar results were reported m of row, 8 in each 10 m of row, to 32 in each 10 m
for purple nutsedge and johnsongrass (Horowitz of row at three different sites. Cotton lint yield
1973). Losses of 16 or 26 percent of the weed-free decreased 6 to 18 kg ha-1 for each added buffalobur
yield were reported by Keeley and Thullen (1991b) in 10 m of row.
when bermudagrass competed with cotton for 12 or
25 weeks. If cotton was hand-weeded for 8 to 12 Coffee SennaCassia occidentalis
weeks after emergence, yield was not affected and With season-long competition, each coffee senna
was 9 percent greater than cotton hand-weeded for plant in 7.5 m of row reduced cotton yield 9 to 117
only 4 weeks after emergence. The future weed kg ha-1 (Higgins et al. 1986). Each additional week
problem from bermudagrass was affected because of competition from 40 coffee senna m-1 of row
bermudagrass did not produce seed, and rhizome reduced cotton yield 118 kg ha-1 and reduced cotton
production was negligible when weed competition stem height 1.25 cm. If cotton was kept weed free
did not exceed 8 weeks after emergence and weed- for 8 or more weeks after emergence, coffee senna
free periods exceeded 4 weeks (Keeley and Thullen did not grow successfully.
In no-till cotton, 3,600 kg ha-1 of bermudagrass Common CockleburXanthium strumarium
reduced cotton height as early as 5 weeks after
planting, and seed cotton yield was reduced 25 per- The dry weight of common cocklebur increased with
cent (Vencill et al. 1992). When bermudagrass was increasing density up to 16 plants in 15 m of row,
present, soil water content was decreased signifi- whereas seed cotton yield decreased as weed density
cantly up to 15 cm deep. Soil water was not signifi- increased up to 16 in 15 m of row (Snipes et al. 1982).
cantly affected 30 to 60 cm deep. In a subsequent Cotton yield losses ranged from 72 to 115 kg ha-1 for
study by Vencill et al. (1993), cotton height and hand-harvested cotton and 57 to 90 kg ha-1 for
yield were significantly reduced by bermudagrass machine-harvested cotton for each common cockle-
and the effects were magnified by increasing weed bur present in 15 m of row. Cotton stem diameter
density. Soil volumetric water content decreased in and height also decreased with increasing weed
the upper 30 cm of soil with increasing bermuda- competition and both were good indicators of com-
grass density. The critical period for bermudagrass mon cocklebur competition. Common cocklebur
competition was 4 to 7 weeks after planting (Vencill reduced cotton yield when competition was longer
et al. 1993). than 4 weeks after emergence in 2 years and 2 weeks
after emergence in a third year (Snipes et al. 1987).
Black NightshadeSolanum nigrum Cotton yield was not affected when it was free of
Black nightshade that competed with cotton for common cocklebur competition for 8 to 10 weeks.
the full season reduced yield 60 to 100 percent. If When cotton was grown less than 60 cm from a
cotton was cultivated about 3 weeks after emer- common cocklebur plant, it was shorter, had less
gence and kept free of black nightshade for the leaf area, and had lower leaf, stem, boll, and bio-
rest of the season, yield was not reduced. The mass dry weight than cotton grown more than 60 cm
largest yield reduction (82 to 100 percent) from a common cocklebur (Byrd and Coble 1991a).
occurred in a year when 0.5 to 0.7 inches of rain Differences in leaf area and biomass were greater 13
fell within 10 days after cotton planting (Keeley weeks after planting, and 15 weeks after planting,
and Thullen 1989a). The importance of regular cotton leaf area and biomass were reduced 11 and
control of black nightshade is emphasized by the 15 percent. When one common cocklebur grew in
fact that fields had to be kept free of black night- 2.1 m of row for 27 weeks after planting, cotton
shade seed production for more than 5 years if yield was reduced 31 percent. Even weed plants 99
populations were to be reduced to a level that did cm from the cotton row reduced cotton yield. Cotton
not affect cotton yield. also competes and common cocklebur grown alone
produced 67 percent more biomass than did cotton
BuffaloburSolanum rostratum grown alone (Byrd and Coble 1991a).
The dry weight of buffalobur increased by 0.06 to
Devils-clawProboscidea louisianica
0.3 kg per plot for each added weed in 10 m of row
(Rushing et al. 1985a). The threshold density for Random densities of 5.5 +- 1.1 devils-claw plants
cotton yield loss varied from 2 buffalobur in each 10 m-2 reduce cotton lint yield 41 kg ha-1 or about 5 per-
The Effect of Weed Density 41

cent for each week the weed was present. Interfer- hogpotato m-2 reduced cotton height after full-
ence of 4, 8, or 12 weeds in each 10 m of row season competition by 14 to 44 percent. The weeds
decreased cotton yield by 22, 49, or 56 kg ha-1, dry weight was reduced 54 percent by full-season
respectively, for each week of interference (Riffle et competition from cotton whose lint yield was
al. 1989). Devils-claw has a deep taproot similar to reduced 31 to 98 percent after full-season competi-
cotton and probably competes primarily for water tion (Castner et al. 1989). Interference during the
and nutrients (Mercer et al. 1987). As devils-claw first 7 weeks of cotton growth reduced lint yield 40
density doubled from 1 to 32 plants 10 m-1 of cotton percent, but if the weed emerged after 7 weeks of
row, cotton yield decreased between 84 and 146 kg crop growth, it had no affect on cotton yield. As is
ha-1. Maximum yield losses ranged from 59 to 74 the case with the deep-rooted perennial devils-claw,
percent over three sites (Mercer et al. 1987). In sup- competition for water is important. Hogpotato uses
port of the hypothesis that devils-claw competes pri- soil water mainly from below 120 cm, while cotton
marily for water, Riffle et al. (1990) demonstrated uses water in the upper 75 cm of soil.
greater water depletion early in the cotton-growing
Ivyleaf MorninggloryIpomoea hederacea
season, which was the time of the weeds most rapid
growth. When cotton was grown alone, the greatest Ivyleaf morningglory planted in California from
water demand was late in the season during peak April through August began to emerge about 1 week
bloom and early boll formation. The vigor of after planting. When the weeds density was one
devils-claw competition and the interaction with plant in 2 m of row and it was present from early
water are illustrated by the finding that in one year, April or May for the entire season, the competition
yield was reduced 96 percent, whereas it was was so severe that the entire cotton crop was lost
reduced only 46 percent in a second year with high- (Keeley et al. 1986). June planting reduced cotton
er rainfall. yield only 11 percent and later plantings had no
effect. The April through July plantings began flow-
Hemp SesbaniaSesbania exaltata
ering within 7 weeks of planting, and viable seed
Hemp sesbania has been studied more as a competi- was collected as soon as 9 weeks after planting. In
tor in soybeans and rice than in cotton. Densities of an Oklahoma study (Wood et al. 1999), cotton yield
1, 2, 5, or 10 plants in 13.3 m of row reduced cotton reductions from 1 weed 10 m-1 of row ranged from
yield 19, 25, 45, or 53 percent, respectively (Bryson 31 to 36 kg ha-1 at one location and from 35 to 36 kg
1987). Cotton plant density, seedling vigor, and the ha-1 at a second location. Lint yield reduction for
number of white blooms per hectare did not differ each weed in 10 m of row ranged from 3.8 to near-
among hemp sesbania densities 75 days after plant- ly 7 percent at the two locations. The weed did not
ing. The weeds height was equal to or greater than affect harvest efficiency at either location.
cottons 55 to 65 days after planting, which suggests
JimsonweedDatura stramonium
that light competition may play a role. Light pene-
tration 1 m above the cotton canopy was 36 percent Variation in rainfall between geographic regions
less than in weed-free plots when hemp sesbania affected jimsonweed competition in cotton but not
density was equal to or greater than 5 per hectare. in soybeans. Jimsonweeds competitive ability was
However, light was only reduced at the soil surface reduced in dry years, but it was always more com-
at the maximum density studied (10 plants in 13.3 m petitive in the less-competitive cotton than in soy-
of row) (Bryson 1987). Although early season vigor beans (Oliver et al. 1991).
and growth of cotton seedlings up to 28 days after
JohnsongrassSorghum halepense
planting was not affected by as many as 3 hemp ses-
bania in 1 m of row (32,000 ha-1), white cotton When johnsongrass competed for the full season, a
blooms and cotton yield were reduced when hemp minimum density of 2 plants in 9.8 m of row was
sesbania was not removed by 70 days after planting required to reduce cotton yield, and yield decline
(Bryson 1990). increased rapidly as density increased (Bridges and
Chandler 1987). The yield loss was 4, 14, 40, 65,
HogpotatoHoffmanseggia glauca
or 70 percent for johnsongrass densities of 2, 4, 8,
Hogpotato is a native perennial, semiprostrate plant 16, or 32 plants 9.8 m-1 of cotton row. Cotton yield
in the southwestern United States, and it rarely loss was proportional to its yield potential at the
grows more than 30 cm tall. As many as 105 +- 21 median densities of 4 and 8 plants m-1 of row, but
42 Chapter 5

not above or below those densities. Rhizome john- Polygonum spp.

songrass was a more vigorous competitor than
The interference of three members of the polygo-
plants from seed. Three to 4 weeks of competition
naceae [ladysthumb (Askew and Wilcut 2002a),
from rhizome johnsongrass decreased cotton yield,
pale smartweed (Askew and Wilcut 2002b), and
but 6 weeks were required before significant yield
Pennsylvania smartweed (Askew and Wilcut
reductions were obtained from seedling john-
2002c)] has been studied in North Carolina. Each
songrass competition (Bridges and Chandler
remained shorter than cotton for 70 to 80 days after
1987). If cotton was maintained weed free for 4
cotton planting. Each grew taller than cotton by har-
weeks after emergence, yield loss in one year was
vest and produced significant dry biomass by cotton
prevented, but it was not in the second year of the
harvest. However, cotton significantly reduced the
study. When cotton cultivars that were 66, 122, or
biomass of each weed by about four times. The
168 cm tall at maturity competed with 4 or 6 john-
hyperbolic function accurately described the rela-
songrass plants 6 m-1 of row with cotton planted in
tionship between each weeds density and cotton
rows 1 m apart, there was no effect of cultivar on
yield loss. In general, cotton yield loss decreased in
competition (Bridges and Chandler 1988). Yield of
the range of 0.7 to 1.3 kg ha-1 for each gram increase
all cultivars declined as johnsongrass density
in weed dry biomass m-1 of crop row.
increased. In the absence of johnsongrass, the yield
of the cultivars did not differ.
PigweedsAmaranthus spp.
In California a weed-free period of 9 weeks after
emergence was required to prevent cotton yield loss The interference of four pigweed species has been
(Keeley and Thullen 1989b). Competition for 6, 9, studied in cotton. Four field experiments were done
12, or 25 weeks after emergence reduced cotton in Oklahoma to determine the effect of full-season
yield 20, 60, 80, or 90 percent. When johnsongrass competition of Palmer amaranth (Rowland et al.
was removed 3 weeks after emergence and the plots 1999). For densities up to 12 weeds 10 m-1 of row,
were kept weed free for the rest of the season, cot- each increase of 1 weed reduced lint yield 58 to 112
ton yield was not reduced. Cotton in plots that were kg ha-1 (roughly 6 to 11 percent) depending on loca-
kept weed free for 3 or 6 weeks (after which weeds tion and year. Lint yield versus end-of-season weed
were allowed to grow) lost 19 or 11 percent of the volume per unit area was linear for almost all loca-
weed-free yield. tions and years. For each m3 increase in weed vol-
The effect on picker versus stripper harvest effi- ume, cotton lint yield decreased 1.5 to 2.3 percent.
ciency of johnsongrass densities of 3, 4, 5, 8, or 15 Lint yield versus end-of-season weed biomass fit a
plants 15 m-1 of row was evaluated in Oklahoma. linear model. Lint yield decreased 5 to 9 percent for
With three or fewer plants in one year and four or each additional kg of Palmer amaranth biomass in a
fewer in a second year, harvest efficiencies were 4.9 plot. No consistent effects on fiber properties were
to 7.6 percent higher for stripper than for picker har- found (Rowland et al. 1999). Smith et al. (2000)
vest. With four or more in one year and five or more found that Palmer amaranth affected lint and seed
in the second, differences in harvest efficiency yield only when weed density was 3,260 ha-1 (the
between the two machine methods were not signifi- highest density studied). Mechanical harvest effi-
cant (Wood et al. 2002). For both years, cotton lint ciency was affected by the highest and lower weed
yield was reduced nearly the same amount (3.5 to densities even though most weed material was dis-
5.5 percent) for the two methods for each john- carded in the field. Palmer amaranth did not affect
songrass plant in 15 m of row. seed moisture content, ginning time, fiber quality, or
the percentage of cleaned lint (Smith et al. 2000).
Noogoora Bur and Fierce Thornapple
When Palmer amaranth density ranged from 0 to 10
Xanthium occidentale and Datura ferox
plants in 9.1 m of row, cotton canopy volume was
Cotton lint yield was reduced 36 and 12 percent and decreased 45 percent 10 weeks after emergence and
the maximum distance of influence was 1.7 m for biomass declined 50 percent 8 weeks after emer-
noogoora bur and fierce thornapple. Noogoora bur gence (Morgan et al. 2001). Similar to the work of
was the more vigorous competitor. Its threshold Rowland et al. (1999), cotton yield decreased linear-
was one weed in 195 m of cotton row and for fierce ly from 13 to 54 percent with 1 to 10 Palmer ama-
thornapple it was one weed in 73 m of row (Charles ranth 9.1 m-1 of row. There was no effect on cotton
et al. 1998). lint properties.
The Effect of Weed Density 43

Smooth pigweed affected cotton by reducing tive (91 percent reduction) in reducing nutsedge
plant water status early in the season and by shading tubers over 3 years.
late in the season (Stuart et al. 1984). With a smooth Yellow nutsedge and johnsongrass had higher
pigweed density of 2.5 m-2, cottons leaf water height, biomass, leaf area, growth rate, and photo-
potential (0.53 MPa) and turgor pressure (0.21 Mpa) synthetic efficiency than cotton or purple nutsedge
reductions were significant. Late in the season, pho- (Holt and Orcutt 1991). Cotton had greater leafiness
tosynthetic photon flux to the cotton canopy at noon and canopy closure than the weeds over 10 weeks
was reduced as much as 90 percent. Smooth pig- but the weeds dominated because they had better
weed was able to maintain a higher water potential overall resource use and production efficiency. Lin-
and turgor pressure than cotton. The weed was able ear correlation analysis indicated that most growth
to extract water from lower in the soil profile, and it variables were significantly correlated with aggres-
had higher diffusive resistance and a lower transpi- sivity; a synonym for competitiveness. Of the 12
ration loss. growth variables studied, 4leaf growth rate,
One redroot pigweed 15 kg ha-1 of row reduced height, relative growth rate, and the growth rate of
cotton yield 21 to 38 kg ha-1 in a study with densi- the initial propagule (seedling or rhizome bud)
ties up to 32 weeds in 15 m of row (Buchanan et al. were best correlated with agressivity. The ability to
1980). Sicklepod was regarded as slightly more capture light and establish a rapidly growing
competitive than redroot pigweed. seedling early was the best predictor of competitive
Full-season interference from 64 tumble pigweed success (Holt and Orcutt 1991).
in 10 m of cotton row reduced lint yield 8 to 11 kg The vigor of competition from yellow nutsedge is
ha-1 for each added weed in 10 m of row. The dry illustrated well in work by Keeley and Thullen
weight of tumble pigweed increased 0.15 to 0.4 kg (1983) who showed that hoeing cotton from 4 to 12
per plot for each added weed in 10 m of row (Rush- weeks after emergence reduced the population of
ing et al. 1985b). The threshold density for lint yield yellow nutsedge present at harvest by 67 to 87 per-
reduction ranged from 4 to 16 tumble pigweeds 10 cent. Over six locations, a yellow-nutsedge-free
m-1 of row. Cotton height was reduced, but only period of 4 to 12 weeks was required to avoid cotton
when the weed density was 32 or 64 10 m-1. yield reduction. All levels of nutsedge control (from
2 to 12 weeks after emergence) reduced the number
Purple and Yellow NutsedgeCyperus rotundus
of nutsedge shoots at harvest and cotton yield loss.
and C. esculentus
Purple nutsedge has a greater leaf area and dry
Keeley (1987) wrote a complete review (93 cita- weight than three other weeds (large crabgrass,
tions) of interference reactions of purple and yellow prickly sida, and velvetleaf) that commonly infest
nutsedge in eight agronomic and nine horticultural cotton. Among these weeds, purple nutsedge was
crops. His review demonstrated that both weeds the most and prickly sida the least competitive
reduce crop yield and both are effective competitors. (Elmore et al. 1983) in competition with cotton and
The articles conclusion is similar to the plea herein, with each other.
a lot is known about what happens when specific
SicklepodCassia obtusifolia
weeds compete with crops. Good descriptions of
what happens are readily available. Keeley (1987) Full-season competition of 4, 12, or 32 sicklepod 15
says perhaps (a weak word) it is time to move on m-1 of row with 5, 10, or 20 cotton plants m-1 of row
to determine why, when, and under what condi- using conventional cultural practices showed no
tions specific crop-nutsedge interactions occur. His effect of cotton density on competitive effects of
plea was, one assumes, heard but not heeded by sicklepod. Cotton yield was inversely related to the
many. Keeley et al. (1979) also showed that crop- weeds density (Street et al. 1981). When sicklepod
ping systems affect yellow nutsedge populations. was combined with redroot pigweed and smooth
Two years of alfalfa or double cropping barley (with pigweed in all possible combinations of 1, 2, 4, 8,
appropriate herbicides) followed by corn that pre- and 16 weeds of each species in 7.5 m of row, seed
ceded cotton, reduced yellow nutsedge tubers by 96 cotton yield declined as a quadratic function of
percent. Two years of chemical fallow (with increasing weed density (Street et al. 1985). Sickle-
glyphosate) following barley and preceding cotton pod was more competitive than either pigweed
removed 98 percent of yellow nutsedge tubers. Con- species. With one pigweed in 7.5 m of row, cotton
tinuous cotton treated with MSMA was also effec- yield decreased 26 kg ha-1. With one sicklepod in 7.5
44 Chapter 5

m of row, yield declined 85 kg ha-1. When the weeds maturing cultivar most. The competitive effect of
were present together with two plants in 7.5 m of spurred anoda and velvetleaf (see velvetleaf studies
row, yield decreased 110 kg ha-1. At low weed den- below) increased with time of competition (Patter-
sity (less than 4 in 7.5 m-1 of row), the competitive son and Highsmith 1989). Growth reduction in cot-
effect of sicklepod and either pigweed was additive. ton was associated with reduced leaf area duration
At higher densities, competition was not additive and the effect was more severe in a dry year.
because of intraspecific competition among the Drought, when imposed, did not affect the relative
weeds (Street et al. 1985). Buchanan et al. (1980) competitive ability of the two weeds or cotton.
found that one sicklepod 15 m-1 of row reduced cot- Intraspecific competition was more severe on cotton
ton yield 34 to 43 kg ha-1. than intraspecific competition.
Silverleaf NightshadeSolanum elaeagnifolium Tropic CrotonCroton glandulosus
This perennial weed with sharp spines is a vigorous Cotton height decreased with increasing weed den-
competitor. It is assumed that competition in cotton sity 10 weeks after planting, but weed height was
is primarily for water because irrigated cotton com- not affected. Cotton lint yield decreased linearly at 2
petes much better than dryland cotton (Green et al. kg ha-1 with each gram increase in tropic croton den-
1987). Cotton height decreased with 4 or more sil- sity m-1 of crop row (Askew and Wilcut 2001). Trop-
verleaf nightshade in 10 m of row. There is no evi- ic croton was less competitive with cotton than other
dence of intraspecific competition and the predicted weeds studied by Askew and Wilcut (2002a, b, c)
cotton yield loss is 1.5 percent per weed in 10 m of but was regarded as an important threat.
row. The weed also interfered with mechanical har- VelvetleafAbutilon theophrasti
vest when density was 16 or 32 10 m-1. When cotton
was grown with silverleaf nightshade, water loss Velvetleaf is a good competitor when soil water is
was greater in the lower parts of the soil profile early adequate because it transpires a lot of water. Light
in the season than when cotton was grown alone interacts with water because solar radiation is the dri-
(Green et al. 1988). The weeds effect on cotton ver of plant water use (Salisbury and Chandler 1993).
yield, height, and boll size was directly correlated In this greenhouse study, velvetleaf used significantly
with the amount of soil water available. There was a more water than cotton when water and light were
negative, linear relationship between cotton lint abundant. Transpiration of both species decreased in
yield and weed biomass. Each 1 kg of weed biomass dry soil. Velvetleaf reduced transpiration losses via
10 m-1 of row of established stands of silverleaf leaf abcission, but cotton leaves did not abcise. Both
nightshade decreased cotton yield 9 percent for 1- species transpired less when shaded, and shading
year-old weeds and 21 percent for 2-year-old weeds. reduced competitive stress in dry soil.
Each stem of the weed in 10 m of row reduced cot- Literature Cited
ton yield about 0.3 percent (Smith et al. 1990).
Askew, S. D., and J. W. Wilcut. 2002a. Ladysthumb
Spotted SpurgeEuphorbia maculata interference and seed production in cotton. Weed
Sci. 50:326332.
Spotted spurge densities of 5, 10, or 50 plants m-1 of
. 2002b. Pale smartweed interference and ach-
row reduced cotton yield 47, 57, or 85 percent,
ene production in cotton. Weed Sci. 50:357363.
respectively, after season-long competition. As the
. 2002c Pennsylvania smartweed interference
weeds density increased, cottons height, leaf area, and achene production in cotton. Weed Sci.
dry weight, and boll number all decreased (Barar- 50:350356.
pour et al. 1994). . 2001. Tropic croton interference in cotton.
Spurred AnodaAnoda cristata Weed Sci. 49:184189.
Bararopour, M. T., R. E. Talbert, and R. E. Frans.
Three cotton cultivars varied in their response to 1994. Spotted spurge (Euphorbia maculata) inter-
spurred anoda competition, but yield of all was ference with cotton (Gossypium hirsutum). Weed
reduced as much as 38 percent by season-long com- Sci. 42:553555.
petition. Cotton yield and the weeds effects varied Bridges, D. C., and J. M. Chandler. 1988. Influence of
due to highly variable weather during the 3 years of cultivar height on competitiveness of cotton
the study (Chandler and Meredith 1983). Early- (Gossypium hirsutum) with johnsongrass (Sorghum
season competition reduced the yield of the early halepense). Weed Sci. 36:616620.
The Effect of Weed Density 45

. 1987. Influence of johnsongrass (Sorghum Hobhouse, H. 1985. Seeds of Change: five plants that
halepense) density and period of competition on transformed mankind. Harper and Row, New York,
cotton yield. Weed Sci. 35:6367. NY. 252 pp.
Bryson, C. T. 1990. Interference and critical time of Holt, J. S., and D. R. Orcutt. 1991. Functional rela-
removal of hemp sesbania (Sesbania exaltata) in tionships of growth and competitiveness in perenni-
cotton (Gossypium hirsutum). Weed Sci. 4:833837. al weeds and cotton (Gossypium hirsutum). Weed
. 1987. Interference of hemp sesbania (Sesba- Sci. 39:575584.
nia exaltata) with cotton (Gossypium hirsutum). Horowitz, M. 1973. Competitive effects of Cynodon
Weed Sci. 35:314318. dactylon, Sorghum halepense, and Cyperus rotun-
Buchanan, G. A., R. H. Crowley, J. E. Street, and J. A. dus on cotton and mustard. Exp. Agric. 9:263273.
McGuire. 1980. Competition of sicklepod (Cassia Keeley, P. E. 1987. Interference and interaction of
obtusifolia) and redroot pigweed (Amaranthus purple and yellow nutsedges (Cyperus rotundus and
retroflexus) with cotton (Gossypium hirsutum). C. esculentus) with crops. Weed Technol. 1:7481.
Weed Sci. 28:258262. Keeley, P. E., and R. J. Thullen. 1991a. Growth and
Byrd, J. D., Jr., and H. D. Coble. 1991a. Interference interaction of barnyardgrass (Echinochloa crus-
of common cocklebur (Xanthium strumarium) and galli) with cotton (Gossypium hirsutum). Weed Sci.
cotton (Gossypium hirsutum). Weed Technol. 39:369375.
5:270278. . 1991b. Growth and interaction of bermuda-
. 1991b. Interference of selected weeds in cot- grass (Cynodon dactylon) with cotton (Gossypium
ton. Weed Technol. 5:263269. hirsutum). Weed Sci. 39:570574.
Castner, E. P. , D. S. Murray, N. M. Hackett, L. M. . 1989a. Growth and competition of black
Verhalen, D. L. Weeks, and J. F. Stone. 1989. Inter- nightshade (Solanum nigrum) and Palmer amaranth
ference of hogpotato (Hoffmanseggia glauca) with (Amaranthus palmeri) with cotton (Gossypium hir-
cotton (Gossypium hirsutum). Weed Sci. sutum). Weed Sci. 37:326334.
37:688694. . 1989b. Growth and interaction of john-
Chandler, J. M., and W. R. Meredith, Jr. 1983. Yields songrass (Sorghum halepense) with cotton (Gossyp-
of three cotton (Gossypium hirsutum) cultivars as ium hirsutum). Weed Sci. 37:339394.
influenced by spurred anoda (Anoda cristata) com- . 1983. Influence of yellow nutsedge (Cyperus
petition. Weed Sci. 31:303307. esculentus)-free periods on yield of cotton (Gossyp-
Charles, G. W., R. D. Murison, and S. Harden. ium hirsutum). Weed Sci. 31:803807.
1998. Competition of Noogoora bur (Xanthium Keeley, P. E., R. J. Thullen, and C. H. Carter. 1986.
occidentale) and fierce thornapple (Datura Influence of planting date on growth of ivyleaf
ferox)with cotton (Gossypium hirsutum). Weed morningglory (Ipomoea hederacea) in cotton
Sci. 48:442446. (Gossypium hirsutum). Weed Sci. 34:906910.
Colvin, D. L., M. G. Patterson, and S. H. Crawford. Keeley, P. E., R. J. Thullen, J. H. Miller, and C. H.
1992. Cotton (Gossypium hirsutum) yield response Carter. 1979. Comparison of four cropping systems
to cultivation, timing and frequency. Weed Technol. for yellow nutsedge (Cyperus esculentus) control.
6:3135. Weed Sci. 27:463467.
Elmore, C. D., M. A. Brown, and E. P. Flint. 1983. Kempen, H. M. 1984. Cotton production losses from
Early interference between cotton (Gossypium hir- weed competition in Kern County: A three year
sutum) and four weed species. Weed Sci. evaluation. Proc,. West Soc. Weed Sci. 37:4751.
31:200207. Klingaman, T. E., and L. R. Oliver. 1994. Influence of
Green, J. D., D. S. Murray, and J. F. Stone. 1988. Soil cotton (Gossypium hirsutum) and soybean (Glycine
water relations of silverleaf nightshade (Solanum max) planting date on weed interference. Weed Sci.
elaeagnifolium) with cotton (Gossypium hirsutum). 42:6165.
Weed Sci. 36:740746. Mercer, K. L., D. S. Murray, and L. M. Verhalen.
Green, J. D., D. S. Murray, and L. M. Verhalen. 1987. 1987. Interference of unicorn-plant (Proboscidea
Full-season interference of silverleaf nightshade louisianica) with cotton (Gossypium hirsutum).
(Solanum elaeagnifolium) with cotton (Gossypium Weed Sci. 35:807812.
hirsutum). Weed Sci. 35:813818. Miller, J. H., L. M. Carter, and C. Carter. 1983. Weed
Higgins, J. M., R. H. Walker, and T. Whitwell. 1986. management in cotton (Gossypium hirsutum) grown
Coffee senna (Cassia occidentalis) competition in two row spacings. Weed Sci. 31:236241.
with cotton (Gossypium hirsutum). Weed Sci. Morgan, G. D., P. A. Baumann, and J. M. Chandler. 2001.
34:5256. Competitive impact of Palmer amaranth (Amaranthus
46 Chapter 5

palmeri) on cotton (Gossypium hirsutum) development hirsutum) densities on competitiveness of pigweed

and yield. Weed Technol. 15:408412. (Amaranthus spp.) And sicklepod (Cassia obtusifo-
Oliver, L. R., J. M. Chandler, and G. A. Buchanan. lia). Weed Sci. 29:253261.
1991. Influence of geographic region on jimson- Street, J. E., C. E. Snipes, J. A. McGuire, and G. A.
weed (Datura stramonium) interference in soybeans Buchanan. 1985. Competition of a binary weed sys-
(Glycine max) and cotton (Gossypium hirsutum). tem with cotton (Gossypium hirsutum). Weed Sci.
Weed Sci. 39:585589. 33:807809.
Patterson, D. T., and M. T. Highsmith. 1989. Competi- Stuart, B. L., S. K. Harrison, J. R. Abernathy, D. R.
tion of spurred anoda (Anoda cristata) and vel- Krieg, and C. W. Wendt. 1984. The response of cot-
vetleaf (Abutilon theophrasti) with cotton ton (Gossypium hirsutum) water relations to smooth
(Gossypium hirsutum) during simulated drought pigweed (Amaranthus hybridus) competition. Weed
and recovery. Weed Sci. 37:658664. Sci. 32:126132.
Riffle, M. S., D. S. Murray, J. F. Stone, and D. L. Vencill, W. K., L. J. Giraudo, and G. W. Langdale.
Weeks. 1990. Soil-water relations and interference 1992. Response of cotton (Gossypium hirsutum) to
between devils claw (Proboscidea louisianica) and coastal bermudagrass (Cynodon dactylon) density
cotton (Gossypium hirsutum). Weed Sci. 38:3944. in a no-tillage system. Weed Sci. 40:455459.
Riffle, M. S., D. S. Murray, L. M. Verhalen, and D. L. . 1993. Soil moisture relations and critical
Weeks. 1989. Duration and intensity of unicorn-plant period of Cynodon dactylon (L.) Pers. (Coastal
(Proboscidea louisianica) interference with cotton bermudagrass) competition in conservation tillage
(Gossypium hirsutum). Weed Technol. 3:313316. cotton (Gossypium hirsutum L.) Weed Res.
Rowland, M. W., D. S. Murray, and L. M. Verhalen. 33:8996.
1999, Full-season Palmer amaranth (Amaranthus Wood, M. L., D. S. Murray, J. C. Banks, L. M. Ver-
palmeri) interference with cotton (Gossypium hir- halen, R. B. Westerman, and K. B. Anderson. 2002.
sutum). Weed Sci. 47:305309. Johnsongrass (Sorghum halepense) density effects
Rushing, D. W., D. S. Murray, and L. M. Verhalen. on cotton (Gossypium hirsutum) harvest and eco-
1985a. Weed interference with cotton (Gossypium nomic value. Weed Technol. 16:495501.
hirsutum). I. Buffalobur (Solanum rostratum). Weed Wood, M. L., D. S. Murray, R. B. Westerman, L. M.
Sci. 33:810814. Verhalen, and P. L. Claypool. 1999. Full-season
. 1985b. Weed interference with cotton interference of Ipomoea hederacea with Gossypium
(Gossypium hirsutum). II. Tumble pigweed (Ama- hirsutum. Weed Sci. 47:693696.
ranthus albus). Weed Sci. 33:815818.
Salisbury, C. D., and J. M. Chandler. 1993. Interaction OILSEED CROPS
of cotton (Gossypium hirsutum) and velvetleaf
(Abutilon theophrasti) plants for water is affected FlaxLinum usitatissimum
by their interaction for light. Weed Sci. 41:6974. Greenhouse studies suggested that dog mustard was
Smith, B. S., J. A. Pawlak, D. S, Murray, L. M. Ver-
less competitive than wheat but similar to flax.
halen, and J. D. Green. 1990. Interference from
Competition from both crops reduced the leaf area,
established stands of silverleaf nightshade
shoot dry weight, height, and seed production of the
(Solanum elaeagnifolium) on cotton (Gossypium
weed compared to its growth on summer fallow
hirsutum)lint yield. Weed Sci. 38:129133.
Smith, D. T., R. V. Baker, and G. L. Steele. 2000.
land. Wall (1997) concluded that dog mustard was
Palmer amaranth (Amaranthus palmeri)impacts on not a vigorous competitor with wheat or flax.
yield, harvesting, and ginning in dryland cotton Literature Cited
(Gossypium hirsutum). Weed Sci. 14:122126.
Snipes, C. E., G. A. Buchanan, J. E. Street, and J. A. Wall, D. 1997. Dog mustard (Erucastrum gallicum)
McGuire. 1982. Competition of common cocklebur response to crop competition. Weed Sci.
(Xanthium pensylvanicum) with cotton (Gossypium 45:397403.
hirsutum). Weed Sci. 30: 553556. Rapeseed = CanolaBrassica napus L.
Snipes, C. E., J. E. Street, and R. H. Walker. 1987.
Interference periods of common cocklebur (Xanthi- The average annual loss in rapeseed yield due to
um strumarium) with cotton (Gossypium hirsutum). infestation with perennial sowthistle was estimated
Weed Sci. 35:529532. to be 9.4 million kg in Saskatchewan and 6.1 million
Street, J. E., G. A. Buchanan, R. H. Crowley, and J. A. kg in Manitoba (Peschken et al. 1983). The weed
McGuire. 1981. Influence of cotton (Gossypium was present in 39 percent of the rapeseed fields
The Effect of Weed Density 47

surveyed but because it existed in patches, it actual- 328 weeds m-2. Predictions based on relative dry
ly infested only about 7 percent of the hectares weight of weeds and crop (weed dry weight/crop +
surveyed. weed dry weight) in December (oilseed rape is a
Wild mustard and common lambsquarters inter- winter crop in the UK) were somewhat less vari-
fered with rapeseed growth early in the season and able than those based only on weed density. In this
caused significant reductions in dry weight by June case, a 5 percent yield loss was caused by 1.4 to
each year. When weed density varied from 20 to 80 10.6 percent range of relative dry weight. Varia-
plants m-2, wild mustard reduced rapeseed grain yield tions in yield loss were caused by variation in
19 to 77 percent, whereas common lambsquarters oilseed rape and common chickweed competitive-
reduced yield only 20 to 25 percent (Blackshaw et al. ness related to weather differences between sites
1987). Both weeds produced abundant seed and, if and years and the 8 to 10 months between planting
not controlled, added large quantities of seed to the and harvest. Lutman et al. (2000) reported that
soil seedbank. As few as 4 wild radish m-2 that despite the variations, there were indications that
emerged with canola reduced canola yield 9 to 11 the greater the crop dry weight was in December,
percent and 64 wild radish m-2 reduced yield 77 to 91 the lower the final crop yield loss. Weed competi-
percent (Blackshaw et al. 2002). Wild radish interfer- tion was not affected by crop density between 44
ence was influenced greatly by its time of emergence and 113 plants m-2 apparently due to the compen-
relative to canola. If it emerged 10 weeks after canola, satory ability of the lowest density.
it had no measurable effect but still produced seed in Canola yield is not affected significantly by usual
most years. Its effect decreased as the time increased. canola planting density. When tartary buckwheat
Rapeseed was less competitive than rye, wheat, or was the competing weed, canola planted at 200
barley in competition with quackgrass in studies in plants m-2 was able to reduce the weeds effect com-
Denmark (Melander 1994) but more competitive pared to densities of 50 to 100 plants m-2 (ODono-
than peas. Yield losses for rapeseed were about 35 van 1994). This planting density may not be
percent from densities of 100 quackgrass shoots m-2 economically feasible if other control methods are
in spring. Prevailing climate conditions did not cheaper. However, with 100 tartary buckwheat
affect yield-density relationships in rye, peas, or bar- plants m-2, canola yield was estimated to be 115 g m-2
ley but had significant effects on rapeseed and if canola density was 50 plants m-2, but yield
wheat. Melander (1994) points out that this was increased to 157 g m-2 when canola density was 200
probably due to the stimulation of quackgrass m-2 (ODonovan 1994). In other work, ODonovan
growth and inhibition of rapeseed growth by the (1991) showed that there was little interspecific
cool rainy weather in the spring of one year. Volun- competition from quackgrass until its density was
teer barley severely reduced canola yield, but the greater than 200 shoots m-2. However, 50 to 100
financial losses were partially offset if a grower was quackgrass shoots m-2 reduced canola yield 18 to 32
able to harvest barley as a crop (ODonovan et al. percent, a significant loss. In most cases the hyper-
1988). Barley was at least 1.5 times more competi- bolic model described the data well (ODonovan
tive in canola than wild oat. 1991, ODonovan et al. 1989). In ODonovan et al.
Oilseed rape fruit weight was reduced by 200 (1989), a population of only 1 wheat plant m-2
wild proso millet plants m-2, fruit number and shoot reduced canola yield 1 percent.
weight were inhibited by 400 plants m-2, and 600
weeds reduced height and delayed flowering (Miller Literature Cited
and Callihan 1995). The number and weight of
Blackshaw, R. E., D. Lemerle, R. Mailer, and K. R.
oilseed rape fruits was reduced one-third after 8 Young. 2002. Influence of wild radish on yield and
weeks of interference compared to 4 weeks of wild quality of canola. Weed Sci. 50:344349.
proso millet interference. With 600 weeds m-2, shoot Blackshaw, R. E., G. W. Anderson, and J. Dekker.
weight was reduced 74 percent, fruit number 85 per- 1987. Interference of Sinapis arvensis L. and
cent, and fruit weight 82 percent after 12 weeks of Chenopodium album L. in spring rapeseed (Brassi-
interference. ca napus L.). Weed Res. 27:207214.
Yield losses caused by common chickweed were Lutman, P. J. W., P. Bowerman, G. M. Palmer, and G.
often high but differed greatly among ten experi- P. Whytock. 2000. Prediction of competition
ments in the UK (Lutman et al. 2000). For exam- between oilseed rape and Stellara media. Weed Res.
ple, a 5 percent yield loss could be caused by 1.4 to 40:255269.
48 Chapter 5

Melander, B. 1994. Modelling the effects of Elymus SunflowerHelianthus annuus L.

repens (L.) Gould competition on yield of cereal,
peas, and oilseed rape. Weed Res. 34:99108. Only four competition studies have been done in sun-
Miller, T. W., and R. H. Callihan. 1995. Interference flower since 1980, and all but one have been done in
between triazine-resistant Brassica napus and Pan- Europe. The high competitive ability of sunflower
icum miliaceum. Weed Res. 35:453460. was verified by studies in Italy of sunflower in com-
ODonovan, J. T. 1994. Canola (Brassica rapa) plant petition with common lambsquarters, wild mustard,
density influences tartary buckwheat (Fagopyrum and greater ammi (Onofrio and Tei 1994). Consistent
tataricum) interference, biomass, and seed yield. with many other studies, the hyperbolic model best
Weed Sci. 42:385389. described the relationship between yield and weed
. 1991. Quackgrass (Eltrygia repens) interfer- density. The three broadleaf weeds had competitive
ence in canola (Brassica campestris). Weed Sci. indices between I = 1.08 and 1.75, which supports the
39:397401. high competitive ability of sunflower. The economic
ODonovan, J. T., A. K. Sharma, K. J. Kirkland, and threshold was 4 to 6 plants m-2 for all three broadleaf
E. A. de St. Remy. 1988. Volunteer barley weeds when control was by hoeing. It was 6 wild
(Hordeum vulgare) interference in canola (Brassica mustard when a postemergence herbicide was used.
campestris and B napus). Weed Sci. 36:734739. Six studies were conducted in southern Spain to
ODonovan, J. T., K. J. Kirkland, and A. K. Sharma. derive competitive models and define the economic
1989. Canola yield and profitability as influenced threshold for corn caraway competition in sunflower
by volunteer wheat infestations. Can. J. Plant Sci. (Carranza et al. 1995). Losses ranged from 19 to 56
percent of the weed-free yield. Consistent with sev-
Peschken, D. P., A. G. Thomas, and R. F. Wise. 1983.
eral other studies, correlation between percent sun-
Loss in yield of rapeseed (Brassica napus, B.
flower loss and weed density were better than those
campestris) caused by perennial sowthistle
with dry weight. When weeds emerged before mid-
(Sonchus arvensis) in Saskatchewan and Manitoba.
Weed Sci. 31:740744. March, they were about 1.5 times more competitive
than those that emerged later. The economic thresh-
SafflowerCarthamus tinctorius L. old to offset the cost of a shallow tillage that may
have achieved 70 percent control ranged from 2.5
When safflower was planted in 11- and 22-cm rows
weeds m-2 for low-yielding sunflower (1,200 kg
at densities of 10 to 192 plants m-2, its yield and
ha-1) to less than 1 weed m-2 for high-yielding sun-
biomass peaked at a density between 70 to 84 m-2.
flower (2,800 kg ha-1) (Carranza et al. 1995).
Decreasing row spacing slightly improved saf-
Season-long competition by kochia densities of 0.3,
flower competition with green foxtail but increas-
1, 3, or 6 plants m-1 of row decreased sunflower ach-
ing crop density had a greater effect (Blackshaw
ene yield 7, 10, 20, or 27 percent, respectively. Just 21
1993). Safflower competing with 500 green foxtail
weeks of competition after sunflower emergence
m-2 increased biomass and seed production up to
decreased yield 6 percent (Durgan et al. 1990).
100 plants m-2 in one year and up to 156 in a sec-
One of the very few studies of competition of a par-
ond year. At these densities, weedy safflower yield-
asitic weed with any crop was done in Spain with nod-
ed less than weed-free safflower, but its yield was
ding broomrape and sunflower (Castejon-Muoz et al.
three to four times more than at lower densities.
1993). Nodding broomrape attached to sunflowers
Safflower is an effective competitor with green
with 6 to 7 leaves and continued to grow and attach
foxtail. High safflower density reduced green fox-
throughout sunflowers vegetative and flowering
tail biomass up to 72 percent and seed yield up to
stages. Extensive, but subterranean, nodding broom-
85 percent. The competition arose from the dense
rape shoot development was observed mainly at sun-
foliar canopy developed by safflower early in the
flowers early heading stage and sunflowers growth
season and its effective shading of the shorter
was reduced. Early planting increased sunflower yield
and reduced nodding broomrapes detrimental effects.
Literature Cited
Literature Cited
Blackshaw, R. E. 1993. Safflower (Carthamus tincto-
rius) density and row spacing effects on competi- Carranza, P., M. Saavedra, and L. Garcia-Torres.
tion with green foxtail (Setaria viridis). Weed Sci. 1995. Competition between Rodolfia segetum and
41:403408. sunflower. Weed Res. 35:369376.
The Effect of Weed Density 49

Castejon-Muoz, M., F. Romero-Muoz, and L. The effects of crop and weed management sys-
Garcia-Torres. 1993. Effect of planting date of tems on weed populations in a corn-corn-peanut
broomrape (Orobanche cernua Loefl.) infections rotation demonstrated that rotation and weed control
in sunflower (Helianthus annuus L.). Weed Res. were both important (Johnson et al. 1992). Florida
33:171176. beggarweed and yellow nutsedge were the dominant
Durgan, B. R., A. G. Dexter, and S. D. Miller. 1990. weeds in corn when a high-input (intensive) weed
Kochia (Kochia scoparia) interference in sunflower management system was used. If no herbicides were
(Helianthus annuus). Weed Technol. 4:5256. used, Florida pusley dominated, illustrating how
Onofrio, A., and F. Tei. 1994. Competitive ability of weed management simultaneously solves and cre-
threshold levels of three broadleaf weed species in
ates weed problems. The weed management system
sunflower. Weed Res. 34:471480.
in peanut became more difficult because the domi-
nant weeds in corn were controlled successively and
Peanut = GroundnutArachis hypogaea L.
the dominant weed in peanut was the more difficult
Florida beggarweed has been studied more than other to control yellow nutsedge.
weeds in peanut. Buchanan and Hauser (1980) stud- The importance of prompt control of sicklepod
ied the effect of Florida beggarweed and sicklepod populations in a peanut-cotton-corn rotation was
grown with peanut in three row widths with the same emphasized in work by Johnson et al. (1994). Each
in-row planting rate for all row widths. Peanut yield year of sicklepod presence at an initial density of 2
increased and weed growth decreased with decreas- sicklepod 9.1 m-1 of row resulted in exponential
ing row width but row width had no effect on peanut increase in sicklepod seedlings in subsequent years.
quality. Peanut yield decreased with increasing time Of the three crops, corn was the most effective com-
of weed competition. Peanut yield, without weed petitor with sicklepod and the weed produced the
interference, in 20.3 cm rows was 6 to 20 percent fewest seed. When sicklepod was present at what
higher than in 81.2 cm rows. Similarly, peanut yield Johnson et al. (1994) defined as a subeconomic
with weed interference in 20.3 cm rows was 8 to 25 threshold density of 2 sicklepod 9.1 m-1 of row that
percent higher over 3 years than in 81.3 cm rows. In was established in the first year of the study, a 7, 21,
a later study, Hauser et al. (1982) showed that Florida and 20 percent increase in sicklepod population in
beggarweed was more detrimental to peanut yield the next 3 years resulted.
than sicklepod when competition lasted all season. Common ragweed height was not affected by its
Peanut yield decreased 15.8 to 30.2 kg ha-1 for each density or by peanut canopy diameter. The weed
Florida beggarweed m-2 whereas for each sicklepod grew taller than peanut throughout the growing sea-
m-2 yield decreased 6.1 to 22.3 kg ha-1. Each kg of son, indicating competition for light was primary
Florida beggarweed decreased peanut yield 0.15 to (Clewis et al. 2001). The rectangular hyperbola
0.74 kg ha-1, and each kg of sicklepod decreased described the relationship in which common rag-
peanut yield 0.08 to 0.23 kg ha-1. Peanut yield corre- weeds aboveground biomass per plant decreased as
lated best with weed dry weight rather than weed its density increased, but the weeds total biomass
population. Florida beggarweed is a good competitor per meter of crop row increased with weed density.
because it grows above the peanut canopy by 52 days Clewis et al. (2001) concluded that common rag-
after planting (DAP), and by 73 days after planting weed is a very competitive weed in peanut and will
photosynthetic active radiation (PAR) reaching cause serious losses if not controlled.
peanuts was reduced 45 percent (Barbour and Common cocklebur has been identified as a major
Bridges 1995). Sicklepod grew above peanut 42 DAP weed in many crops in the southern United States.
and reduced PAR 41 percent at 79 DAP. Wild poin- With common cocklebur densities of 2, 4, 8, 16, or 32
settia grew above peanut 44 DAP and reduced PAR weeds 8 m-1 of row, yields were reduced 32, 48, 65, or
39 percent at 85 DAP (Bridges et al. 1992). By these 88 percent after full-season interference in one year
criteria, the three weeds ought to be equally competi- and 18, 30, 46, 62, or 75 percent in a second year,
tive with peanut, and in the Barbour and Bridges respectively (Royal et al. 1997). The reduced effect in
(1995) study they were. The distance of influence for the second year was due to that year having above-
the three weeds was Florida beggarweed162 cm, normal rainfall, which either reduced the weeds com-
sicklepod150 cm, and wild poinsettia190 cm. petitive ability or enhanced the crops.
Yield losses within the distance were 26, 27, and 22 Predicted yield losses due to wild poinsettia in
percent, respectively. peanut were 4, 8, 12, 15, 26, 40, or 54 percent for
50 Chapter 5

season-long interference of densities of 1, 2, 4, 8, dal weed control. There was greater early-season
16, or 32 wild poinsettia 9 m-1 of row in Georgia. peanut canopy development when thrips were con-
Losses from the same densities in Florida were trolled, but there was no increase in peanut yield
approximately the same (9, 14, 22, 30, 37, or 41 per- due to thrip control.
cent; Bridges et al. 1992).
After full-season interference from 8, 16, 32, or Literature Cited
64 bristly starbur plants 7.5 m-1 of row, peanut yield Barbour, J. C., and D. C. Bridges. 1995. A model of
was reduced 14, 26, 43, or 50 percent. If peanuts competition for light between peanut (Arachis
were kept weed free for 6 weeks after emergence, hypogaea) and broadleaved weeds. Weed Sci.
the seed yield was reduced not more than 3 percent 43:247257.
(Walker et al. 1989). When peanut density was 72 Bridges, D. C., B. J. Brecke, and J. C. Barbour. 1992.
plants 7.5 m-1 of row, bristly starbur interference for Wild poinsettia (Euphorbia heterophylla) interfer-
2 weeks after emergence reduced seed yield an aver- ence with peanut (Arachis hypogaea). Weed Sci.
age of 4 percent. Yield was reduced 54 percent after 40:3742.
13 weeks of interference. With a bristly starbur den- Buchanan, G. A., and E. W. Hauser. 1980. Influence
sity of 35 plants 7.5 m-1 of row, peanut reduced the of row spacing on competitiveness and yield of
weeds dry weight about 32 percent after 13 weeks peanuts (Arachis hypogaea). Weed Sci. 28:401409.
of interference. Chamblee, R. W., L. Thompson, Jr., and H. D. Coble.
1982. Interference of broadleaf signalgrass
Horsenettle presence for 6 to 8 weeks after emer-
(Brachiaria platyphylla) in peanuts (Arachis
gence did not reduce yield of Spanish runner
hypogaea). Weed Sci. 30:4549.
peanuts, and weed-free maintenance for 2 weeks
Clewis, S. B., S. D. Askew, and J. W. Wilcut. 2001.
after emergence increased yield (Hackett et al.
Common ragweed interference in peanut. Weed Sci.
1987). Linear regression predicted a peanut yield 49:768772.
increase of 69 kg ha-1 for each week of weed main- Hackett, N. M., D. S. Murray, and D. L. Weeks. 1987.
tenance after emergence or a 40 kg ha-1 yield Interference of horsenettle (Solanum carolinense)
decrease for each week of weed interference but with peanuts (Arachis hypogaea). Weed Sci.
only in one year. The work seems to indicate that 35:780784.
horsenettle may not be a major problem. In one year, Hauser, E. W., G. A. Buchanan, R. L. Nichols, and R.
32 weeds in 10 m of row, the highest density in the M. Patterson. 1982. Effects of Florida beggarweed
study (35,200 in an acre), reduced yield, but in a (Desmodium tortuosum) and sicklepod (Cassia
second year, the same density had no effect on yield. obtusifolia) on peanut (Arachis hypogaea) yield.
The interference of two grasses has been reported. Weed Sci. 30:602604.
Full-season interference of a natural infestation of Johnson, W. C., III, J. Cardina, and B. G. Mullinix.
broadleaf signalgrass at 8, 16, or 1,050 plants 10 1992. Crop and weed management effects on weed
m-1 of row reduced peanut seed yield 14, 28 or 69 populations in a short-term corn-corn, peanut rota-
percent. Slightly less than 4 weeds 10 m-1 of row tion. J. Prod. Agric. 5:566570.
reduced yield significantly. If broadleaf signalgrass Johnson, W. C., III, J. Cardina, and B. G. Mullinix, Jr.
was present for 6 weeks or less after emergence, 1994. Dynamics of subeconomic threshold popula-
yield was not affected, but 8 weeks of interference tions of sicklepod (Cassia obtusifolia) in a peanut-
or longer reduced yield (Chamblee et al. 1982). cotton-corn rotation. Weed Sci. 42:364368.
Murdock, E. C., J. A. Alden, and J. E. Toler. 1986.
Peanut is a weaker competitor because full-season
Interactive effects of tobacco thrips control and her-
interference by broadleaf signalgrass reduced
bicides on competition between large crabgrass
peanut forage yield 64 percent, whereas peanut
(Digitaria sanguinalis) and peanuts (Arachis
interference for the full-season reduced the weeds
hypogaea). Weed Sci. 34:896900.
yield only 10 percent. Royal, S. S., B. J. Brecke, and D. L. Colvin. 1997.
One study reported the interaction between Common cocklebur (Xanthium strumarium) inter-
tobacco thrips (Frankliniella fusca Hinds) control ference with peanut (Arachis hypogaea). Weed Sci.
with aldicarb [2-methyl-2-(methylthio)propi- 45:3843.
onaldehyde-O-(methyl-carbamoyl)oxime] and Walker, R. H., L. W. Wells, and J. A. McGuire. 1989.
large crabgrass interference in peanut (Murdock et Bristly starbur (Acanthospermum hispidum) inter-
al. 1986). Thrip control did not affect large crab- ference in peanut (Arachis hypogaea). Weed Sci.
grass dry weight or peanut yield without herbici- 37:196200.
The Effect of Weed Density 51

POTATOSOLANUM TUBEROSUM L. 0.04 and 2 shoots m-2 or 0.0165 and 1.5 g of total dry
A natural stand of grasses including barnyardgrass
Potato cultivars Atlantic and Russet Burbank and
could compete with the potato cultivar Superior for 6
barnyardgrass were more competitive than redroot
to 8 weeks before yield was depressed. On the other
pigweed when the measure was relative competitive
hand, a 2 to 4 week weed-free period was sufficient to
ability (VanGessel and Renner 1990b). In additive
assure no loss of potato yield (Vitolo and Ilnicki
design field studies, 4 redroot pigweed plants or 4
1985). Raby and Binning (1985) affirmed that potato
barnyardgrass plants in each m of row did not reduce
cultivars differ in their competitive ability.
the yield of Atlantic potatoes when the weeds were
A mixture of annual weeds that emerged 1 week
planted between crop rows following hilling 6 to 7
after potatoes and competed for the full season,
weeks after planting. However, a single plant of either
reduced yield an average of 54 percent compared to
weed in a m of row reduced tuber yield 19 to 33 per-
only 16 percent loss when weeds emerged 3 weeks
cent when the weeds were seeded in the potato row
after potatoes (Nelson and Thoreson 1981). With
when potatoes were planted. Aboveground potato bio-
full-season competition, each additional 10 percent
mass was not a consistent predictor of total tuber
of total weed biomass reduced tuber yield 12 per-
yield. The variability of aboveground biomass was not
cent. The reduction in tuber yield was calculated as
due to increases in weed weight or density. For Russet
Burbank and Atlantic potatoes, a single hilling was not
adequate in either of two years for full-season weed
percent reduction in tuber yield from weeds = 47.5 control (VanGessel and Renner 1990a). Early hilling
+ 1.23 (percent of total biomass as weeds) + tended to increase the biomass of C4 compared to C3
0.0045 (days)2 weeds on mineral and muck soil, but it provided ade-
quate weed control in only one year on mineral soil.
In the equation, days represent the number of days Weeds reduced aboveground potato biomass on both
between planting and harvest (Nelson and Thoreson soils, but this was not indicative of tuber yield. Russet
1981). Burbank potato yield was reduced by weeds more
Potato yield decreased with increasing quackgrass than Atlantic on mineral soil when conventional cul-
density and duration of competition. Quackgrass had tural practices (two hillings) were used; the opposite
a greater influence on marketable tuber yield than on was true on muck soil (VanGessel and Renner 1990a).
total yield (Baziramakenga and Leroux 1994). The
Literature Cited
duration of the critical period for weed control varied
with quackgrass density and year. If a 5 percent yield Baziramakenga, R., and G. D. Leroux. 1994. Critical
loss was deemed acceptable, the critical period began period of quackgrass (Eltrygia repens)removal in
15 days after potato emergence at a low level of potatoes (Solanum tuberosum). Weed Sci.
quackgrass interference (35 to 38 g m-2) to approxi- 42:528533.
mately 3 days after emergence at a medium infestation . 1998. Economic and interference threshold
level of 87 to 95 m-2. With a high level of infestation densities of quackgrass (Eltrygia repens) in potato
(135 to 158 g m-2), the critical period began prior to (Solanum tuberosum). Weed Sci. 46:176180.
potato emergence, so one must conclude there was no Nelson, D. C., and M. C. Thoreson. 1981. Competi-
critical period because weed control was required tion between potatoes (Solanum tuberosum) and
weeds. Weed Sci. 29:672677.
from crop emergence on. Weed control was not
Raby, B. J., and L. K. Binning. 1985. Weed competi-
required 23 to 68 days after emergence depending on
tion studies in Russet Burbank and Superior potato
quackgrass density and year. Because the onset of
(Solanum tuberosum) with different management
interference varied less than the end, early control of practices. Proc. N. Central Weed Cont. Conf. 40:4.
quackgrass is required (Baziramaenga and Leroux VanGessel, M. J., and K. A. Renner. 1990a. Effect of
1994). In further studies of quackgrass interference in soil type, hilling time, and weed interference on
potato, Baziramakenga and Leroux (1998) determined potato (Solanum tuberosum) development and
that the dry weight of quackgrass was the best deter- yield. Weed Technol. 4:299305.
minant of potato yield loss. A yield loss of 10 percent . 1990b. Redroot pigweed (Amaranthus
was caused by 25 quackgrass shoots m-2, which was retroflexus) and barnyardgrass (Echinochloa crus-
equivalent to 20 g of total dry biomass. The economic galli) interference in potatoes (Solanum tubero-
threshold for quackgrass in potatoes varied between sum). Weed Sci. 38:338343.
52 Chapter 5

Vitolo, D. B., and R. D. Ilnicki. 1985. Grass competi- Thirty years later, Ni et al. (2000) described near-
tion in white potatoes. Abstr. Weed Sci. Soc. Ameri- ly the same traits that conferred competitive ability:
ca, p. 30. initial biomass, plant growth rate, leaf area index,
RICEORYZA SATIVA L. and biomass at tillering. Biomass at tillering was the
best predictor of competitiveness against weeds in
If the number of people who depend on a crop for a this study, which included newer (different) varieties
major portion of their daily food is the most appro- than those studied by Jennings and Aquino (1968a,
priate measure of importance, then rice is the b, c). The importance of high tillering capacity to
worlds most important crop. The International Rice breeding efforts to maximize the ability of rice cul-
Research Institute has claimed that a third of the tivars to compete with weeds was again emphasized
worlds people rely on rice for 50 percent of their by Estorinos et al. (2002).
daily caloric intake. The world grows 153.8 million A study of the competitiveness of cultivars of
ha of rice annually. The average worldwide yield is upland rice under low-input conditions in the Ivory
3,885 kg ha-1, which gives a worldwide production Coast showed that cultivar competitiveness was cor-
of 598.8 million metric tons, a total production related with root growth at early growth stages and
greater than that of either corn (590.8 million metric with shoot and root growth at later growth stages
tons) or wheat (576.3 million metric tons) (United (Fofana and Rauber 2000).
Nations 2000). Rice is the only major grain crop that Work by Lindquist and Kropff (1996) emphasized
is grown almost exclusively for human food. It is the importance of early leaf area expansion to com-
also the only major grain crop that is grown in stand- petitive ability because of the central role of light
ing water (although it does not have to beupland capture. Their ecophysiological approach predicted
rice is common in South America) and that is eaten that the leaf area index 70 to 75 days after planting
with little additional processing (it is dehulled) after was a good indicator of leaf area expansion rate. The
harvest. model showed that if the early leaf area expansion
It may be related to the crops importance or sim- rate could be increased, barnyardgrass seed produc-
ply to the interests of the scientists, but compared to tion decreased. Therefore, competitive rice cultivars
other crops, the work on rice reported here has more could reduce the need for other weed management
emphasis on exploration of why rather than on what techniques. Detailed study of three rice cultivars
happens in crop-weed competition. In a series of showed that the cultivar that accumulated more bio-
studies, Jennings and Aquino (1968a, b, c) defined mass had a higher leaf area index, a higher specific
varietal traits that made some cultivars more com- leaf area, and, especially in early growth stages, par-
petitive than others. High-tillering, leafy tropical titioned more biomass to leaves was the most com-
indica rices were more competitive when mixed petitive with weeds (Johnson et al. 1998).
with small, erect, sturdy plant types even though In contrast to the preceding studies but not in any
pure stands of the latter always outyielded the for- essential disagreement, Pantone et al. (1992) used
mer (1968a, b). Competition was first observed path analysis to show that the number of panicles per
when plants were 53 to 60 days old, which was 30 plant and florets per panicle were the yield compo-
to 35 days after transplanting. Tall and dwarf culti- nents that determined the responses of fecundity and
vars differed genetically in ways that affected tiller- grain yield to competition. The effects of density on
ing, leaf number, leaf length and angle, and height. percent filled florets and grain weight varied and
Tall genotypes were more competitive under normal were relatively small, suggesting that these things
growth conditions and became relatively more so in were determined primarily by density-independent
response to fertility and close spacing (1968b). The factors.
number of tillers, leaf number, leaf length, leaf area Ahmed and Hogue (1981), in Bangladesh,
index, height, and dry weight were always greater in defined cultivar height as an important characteris-
successful competitors before competition was tic. Yield reduction from weeds increased with
observed (1968c). Leaf length was a critical factor decreasing plant height. It is a logical assumption
because it determined the angle or degree of erect- that this is related to competition for light as
ness and the amount of light the leaf could receive. described by Caton et al. (2001) who used a
Jennings and Aquino (1968c) concluded that any rice:weed model to analyze the effects of the leaf
plant trait that increased size and vigor during early area density (LAD) of weeds, leaf angles, and max-
growth conferred competitive ability. imum height on growth and competition of weeds
The Effect of Weed Density 53

with rice. Short weeds and weeds with conical Nyarko and DeDatta (1993a) studied the interaction
LADs were weakly competitive regardless of other of light and nitrogen when rice competed with itch-
traits. For other weeds, interference with rice was grass and junglerice in the field. Nitrogen availabil-
positively correlated with maximum height, LAD, ity increased the canopy light absorption coefficient
and leaves that were planophile (oriented parallel to and reduced the sunlit leaf area index of rice. When
the ground). Ampong-Nyarko et al. (1992) exam- rice was grown in a growth chamber with low pho-
ined the response of upland rice and three C4 weeds tosynthetically active radiation (PAR), it had higher
(junglerice, goosegrass, and itchgrass) to low light shoot nitrogen concentration than when it was
intensities (150, 250, and 400 mol m-2s-1 of photo- grown at higher PAR. Rices photosynthetic rate was
synthetically active radiation). All three weeds had highly correlated with leaf N content per unit leaf
higher net CO2 exchange rates than rice at all light area. Without nitrogen application, there was no dif-
intensities. The response to CO2 exchange rates was ference in rices dry matter yield at 150 versus 400
greatest when plants were young and gradually mol m-2s-1. The limited response of rice to nitrogen
decreased as plants matured. Itchgrass is an impor- applied to shaded plants and acclimation of rice to
tant weed in the tropics and has superior growth and reduced light could be significant factors in light
carbon assimilation compared to rice under low and and N interaction in rice-weed competition. In fur-
high light intensity. The other two weeds were more ther work, Ampong-Nyarko and DeDatta (1993b)
susceptible to the negative effects of shading. suggested that timing of nitrogen application could
Further evidence of the important role of light in be employed for weed management, although there
rice-weed competition is provided by the work of is no evidence that it has been.
Gibson et al. (2002). Late watergrass seeded with In field studies in Greece (Eleftherohorinos
water-seeded rice was not affected by rice. When the 2002), interference between red rice and two rice
weed was seeded after rice, shading by the crop cultivars began 3 weeks after emergence and was
increased and competition was effective. Gibson et not affected by increasing nitrogen fertility from
al. (2002) proposed that management strategies that 100 to 150 kg ha-1. One cultivar (Thaibonnet) was
delay germination and growth of late watergrass and affected more than the other and its yield was
other Echinochola species might confer a competi- reduced 58 percent by 40 red rice plants m-2 and the
tive advantage to rice and reduce the need for herbi- other (Ariette) was reduced only 46 percent. Red
cide applications. However, the grass is a good rice grew taller than both cultivars 10 weeks after
competitor that can reduce the yield of rice by 18 planting and light competition may have been
percent after only 30 days of competition. In an ear- important.
lier study (Gibson et al. 1999), root competition was Smith was one of the most productive of the weed
identified as the primary mechanism determining scientists who worked on rice. In 1988, Smith iden-
competition between water-seeded rice and late tified the weeds that were most damaging to rice
crabgrass. Gibson et al. (1999) suggested that yield in Arkansas and ranked them. Of the grasses,
researchers should not rely solely on correlations red rice reduced rice yield the most followed by
between shoot traits and competitive ability as evi- barnyardgrass, bearded sprangletop, and broadleaf
dence that competition is primarily for light. Shad- signalgrass. Among the broadleaf/aquatic weeds,
ing by rice had little effect on late watergrass when hemp sesbania reduced rice yield the most followed
it and rice were seeded at the same time. by northern jointvetch, ducksalad, spreading
Ampong-Nyarko and DeDatta (1993a) studied the dayflower, and eclipta. He also reported that barn-
response of four weeds (spiny amaranth, goose- yardgrass, broadleaf signalgrass, and ducksalad
grass, itchgrass, and purple nutsedge) and rice to interfered with rice the most during the early season,
nitrogen. The nitrogen response of two rice cultivars and eclipta, hemp sesbania, northern jointvetch, red
and itchgrass reached a plateau between 69 and 103 rice, and spreading dayflower were more detrimen-
mg N kg-1 of air-dried soil, whereas the other weeds tal from midseason to late season.
continued to respond to increasing amounts of nitro- Pantone and Baker (1991) used reciprocal yield
gen. At the higher nitrogen rates, nitrogen uptake by analysis to study red rice interference. Over 4 years,
weeds was higher than that of rice. If nitrogen was 1 red rice plant reduced rice yield as much as 4 rice
applied at rates that were suboptimal for rice pro- plants of the same cultivar. Rice yield losses from
duction, the competitive ability of rice was reduced red rice interference were 13, 37, 48, or 92 percent
in the presence of each of the four weeds. Ampong- from densities of 4, 16, 25, or 300 red rice plants m-2.
54 Chapter 5

Season-long densities of 1, 2, 5, 10, 20, or 40 red when light is abundant. The addition of phosphate
rice m-2 demonstrated that interference occurred fertilizer stimulates the weeds tillering and final
with as few as 2 red rice m-2 (Kwon et al. 1991b). tiller number is influenced by nitrogen supply.
Ten red rice m-2 reduced total milled and head rice of Barnyardgrass reduced the yield of a semidwarf
the semidwarf cultivar Lemont but did not affect the (Lemont) cultivar more than that of a short-stature
short-statured cultivar Newbonnet. The difference cultivar (Newbonnet) (Stauber et al. 1991). Season-
was due to the shading effect of red rice on the semi- long interference of 20 barnyardgrass plants m-2
dwarf cultivar. When 20 red rice m-2 were grown for reduce yield of the semidwarf cultivar 301 kg ha-1
120 days after rice emergence, the straw weight of and of the short-statured cultivar 257 kg ha-1. The
Lemont was reduced 58 percent and the taller New- importance of proximity is illustrated by the fact
bonnet was reduced 34 percent. Grain yields were that when barnyardgrass was 25 to 50 or 50 to 100
reduced 86 percent in Lemont and 52 percent in cm away from rice plants, there was no effect on rice
Newbonnet. These effects were attributed to the dif- yield. When a barnyardgrass plant group (4 plants in
ference in shoot morphology of the two cultivars 140 cm2 ) was within 25 cm of rice plants, rice yield
and to the vigor of red rice competition (Kwon et al. was reduced up to 21 percent. Perera et al. (1992)
1991a). studied barnyardgrass interference in rice in Sri
Red rice at a density of 5, 108, or 215 plants m-2 Lanka and provided an explanation for the quantita-
reduced rice yield 22, 77, or 82 percent, respective- tive effects observed by others. They used rice
ly, when cultivated rice density was 195 plants m-2 grown in bags sunk in a rice paddy so that roots
(Diarra et al. 1985). Only five red rice m-2 reduced could intermingle with weed roots in one bag, or be
rice grains per panicle by 8 to 18 percent, and 108 or separated from other roots. Thus, they were able to
215 reduced grains per panicle 56 or 70 percent. A calculate the relative importance of shoot and root
medium grain cultivar (Mars) that matured in 138 competition among three rice cultivars with differ-
days in Arkansas competed better than Lebonnet, a ent shoot morphology. Independent of shoot mor-
long-grain cultivar that matured in 126 days. phology, root was always more important than shoot
Ferrero (1996) used a day-degree predictive model competition. Perera et al. (1992) concluded that
for growth of roundleaf mudplantain in competition inhibition of rice root growth led to a reduced capac-
with rice. The weed was allowed to emerge at 7-day ity to take up nutrients from soil and was the most
intervals for 49 days after rice emergence. Consistent important factor in the interference between barn-
with results from other studies on rice and other yardgrass and rice.
crops, the weeds that emerged first were the most Red rice (Diarra et al. 1985) and bearded sprangle-
damaging to yield. The results of the day-degree top (Smith 1983) are more vigorous competitors than
model were consistent in that the weeds that accumu- broadleaf signalgrass. A density of 180 broadleaf sig-
lated (emerged earliest) the greatest number of day nalgrass m-2 reduced rice dry yield a maximum of 48
degrees (403) caused the greatest loss (95 percent). percent 95 days after rice emergence for the cultivar
Echinochloa spp. were much more competitive in Bond. For Mars, a more competitive cultivar (Pantone
direct-seeded than in transplanted rice (Hill et al. and Baker 1991), the maximum yield reduction after
1989). Three Echinochloa spp. m-2 reduced direct- season-long interference was only 21 percent
seeded rice yield 20 percent, but 6.6 weeds were (McGregor et al. 1988a). Each broadleaf signalgrass
required to give the same yield reduction in trans- reduced rice yield by 18 kg ha-1 in 2 years of study in
planted rice. In terms of competitiveness, 25 trans- Arkansas (McGregor et al. 1988b). However, the
planted rice plants m-2 were equal to 300 m-2 in results over 2 years were quite different. In one year,
direct-seeded rice. The regression model developed 50, 100, or 150 weeds m-2 reduced rice dry weight 6
by Hill et al. (1989) showed that total plant stand and weeks after emergence, but in the second year, only
the dependent variable, relative yield, were more use- the highest density (150 weeds m-2 ) reduced rice dry
ful measures of competitive effects than the more weight. McGregor et al. (1988b) suggested the differ-
commonly employed weed density and crop yield. ence between the years was related to the presence in
The importance of early competition as a deter- the second year of barnyardgrass and red rice as addi-
minant of the effect of barnyardgrass on rice is tional competitors. The difference was not related to
emphasized in the work of Kleinig and Noble weather.
(1968). Barnyardgrass grows rapidly and tillers Yields of direct-seeded paddy rice at optimum
abundantly (earlier than rice) early in the season stands of 215 to 270 plants m-2 were reduced 9, 18,
The Effect of Weed Density 55

20, or 36 percent by bearded sprangletop densities tition in rice. Int. Rice Res. Inst. Newsletter 6(3):20
of 11, 22, 54, or 108 plants m-2 (Smith 1983). There Ampong-Nyarko, K., and S. K. DeDatta. 1993a.
was a linear decrease of 21 kg ha-1 for each bearded Effects of light and nitrogen and their interaction
sprangletop m-2. Weed densities of 54 and 108 m-2 on the dynamics of rice-weed competition. Weed
reduced whole milled rice kernel yield and the high- Res. 33:18.
est density (108 m-2) reduced rice seed germination. . 1993b. Effects of nitrogen application on
In a second study, Carey et al. (1994) showed that growth, nitrogen use efficiency and rice-weed inter-
durations of 63, 70, or 130 days after rice emergence action. Weed Res. 33:269276.
led to yield losses of 11, 21, or 50 percent for the Ampong-Nyarko, K., S. K. DeDatta, and M.
Dingkuhn. 1992. Physiological response of rice and
semidwarf cultivar Lemont, and of 11, 13, or 37 per-
weeds to low light intensity at different growth
cent for the conventional cultivar Newbonnet, the
stages. Weed Res. 32:465472.
better competitor because of its height. Competitive
Carey, V. F, III, R. J. Smith, Jr., and R. E. Talbert.
durations of 21 to 56 days after emergence did not
1994. Interference durations of bearded sprangletop
affect yield of either cultivar. (Leptochloa fascicularis) in rice (Oryza sativa).
With the same crop density as the previous study Weed Sci. 42:180183.
(Smith 1983), drill-seeded paddy rice yields were Caton, B. P., T. C. Foin, and J. E. Hill. 1997. Mechanisms
reduced 18 percent by 22 spreading dayflower m-2 of competition for light between rice (Oryza sativa)
competition for 125 to 140 days (full season) (Smith and redstem (Ammania spp.). Weed Sci. 45:269275.
1984). Competition durations up to 80 days did not Caton, B. P., A. M. Mortimer, T. C. Foin, J. E. Hill, K.
affect rice yield. D. Gibson, and A. J. Fischer. 2001. Weed shoot
When rice was planted in a greenhouse with 400 morphology effects on competitiveness for light in
seeds m-2 and redstem was planted at 50 or 100 direct-seeded rice. Weed Res. 41:155163.
seeds m-2 , redstem was taller than rice 45 days after Diarra, A., R. J. Smith, and R. E. Talbert. 1985. Inter-
planting (Caton et al. 1997). By midseason (57 days ference of red rice (Oryza sativa) with rice (O.
after planting), redstem had no observable effects on sativa). Weed Sci. 33:644649.
any plant variable. By final harvest (110 to 118 days Eleftherohorinos, I. G., K. V. Dhima, and I. B. Vasi-
after planting), redstem at both densities reduced lakoglou. 2002. Interference of red rice in Greece.
rice tiller density, panicle density, shoot biomass, Weed Sci. 50:167172.
and grain weight. The weeds effects were only Estorninos, L. E. Jr., D. R. Gealy, and R. E. Talbert.
observed (measurable) after it had grown above the 2002. Growth response of rice (Oryza sativa) and
crop canopy. Thus, the competitive effects were due red rice (O. Sativa) in a replacement series study.
to shading and light competition. Season-long com- Weed Technol. 16:401406.
petition reduced rice yield 31 and 39 percent at the Ferrero, A. 1996. Prediction of Heteranthera reni-
two densities. Caton et al. (1997) classified redstem formis competition with flooded rice using day
degrees. Weed Res. 36:197202.
as the most competitive dicot weed they had studied.
Fofana, B., and R. Rauber. 2000. Weed suppression
Ransom and Oelke (1982) studied the interference
ability of upland rice under low-input conditions in
of common water plantain in wild rice (Zizania palus-
West Africa. Weed Res. 40:271280.
tris). Wild rice cultivars did not differ in their response
Gibson, K. D., A. J. Fischer, T. C. Foin, and J. E. Hill.
to interference and no density of water plantain grown 2002. Implications of delayed Echinochloa spp.
from seed with up to 82 m-2 reduced rice yield. How- Germination and duration of competition for inte-
ever, water plantain established from rootstocks sig- grated weed management in water-seeded rice.
nificantly reduced rice yield at densities as low as 3 Weed Res. 42:351358.
plants m-2. A density of 43 plants m-2 established from Gibson, K. D., T. C. Foin, and J. E. Hill. 1999. The
rootstocks reduced wild rice yield 91 percent. If the relative importance of root and shoot competition
water plantain was removed by 7 weeks after planting, between water-seeded rice and Echinochloa phyllo-
there was no effect on yield but interference for 9 pogon. Weed Res. 39:181190.
weeks or longer reduced yield. The yield component Hill, J. E., S. K. DeDatta, and J. G. Real. 1989.
most affected was panicles per plant. Echinochloa competition in rice: A comparison of
studies from direct-seeded and transplanted flooded
Literature Cited
rice. pp. 115129 in B. A. Auld, R. C. Umali, and
Ahmed, N. U., and M. Z. Hogue. 1981. Plant height S. S. Thifrosomo (ed.) Biotrop. Spec. Pub. No. 39.
as a varietal characteristic in reducing weed compe- Bogor, Indonesia.
56 Chapter 5

Jennings, P. R., and R. C. Aquino. 1968a. Studies on Smith, R. J., Jr. 1983. Competition of bearded spran-
competition in rice. I. Competition in mixtures of gletop (Leptochloa fascicularis) with rice (Oryza
varieties. Evolution 22:119124. sativa). Weed Sci. 31:120123.
. 1968b. Studies on competition in rice. II. . 1984. Competition of spreading dayflower
Competition in segregating populations. Evolution (Commelina diffusa) with rice (Oryza sativa). Weed
22:332336. Sci. 32:116119.
. 1968c. Studies on competition in rice. III. . 1988. Weed thresholds in southern U.S. rice.
The mechanism of competition among phenotypes. Weed Technol. 2:232241.
Evolution 22:529542. Stauber, L. G., R. J. Smith, Jr., and R. E. Talbert.
Johnson, D. E., M. Dingkuhn, M. P. Jones, and M. 1991. Density and spatial interference of barnyard-
C. Mahamane. 1998. The influence of rice plant grass (Echinochloa crus-galli) with rice (Oryza
types on the effect of weed competition on Oryza sativa). Weed Sci. 39:163168.
sativa and Oryza glaberrima. Weed Res. United Nations Food and Agriculture Organization.
38:207216. 2000. Production yearbook 54:7475.
Kleinig, C. R., and J. C. Noble. 1968. Competition
between rice and barnyardgrass (Echinochloa). 1. SORGHUMSORGHUM BICOLOR (L.)
The influence of weed density and nutrient supply MOENCH
in the field. Aust. J. Exp. Agric. and Anim. Husb.
8:358363. Barnyardgrass, large crabgrass, and Texas panicum
Kwon, S. L., R. J. Smith, Jr., and R. E. Talbert. 1991a. were evaluated in the field over 3 years to determine
Interference durations of red rice (Oryza sativa) in the effect of interference duration and weed density
rice (O. sativa). Weed Sci. 39:363368. on sorghum yield (Smith et al. 1990). Linear regres-
. 1991b. Interference of red rice (Oryza sativa) sion predicted a yield loss of 3.6 percent for each
densities in rice (O. sativa). Weed Sci. 39:169174. week of weed interference regardless of year or the
Lindquist, J. L., and M. J. Kropff. 1996. Applications weed species. When grain sorghum was grown in
of an ecophysiological model for irrigated rice 61-cm rows, it was not affected much by full-season
(Oryza sativa)Echniochloa competition. Weed interference from any of the three grass weeds. If
Sci. 44:5256. rows were 91 cm, the effects of interference
McGregor, J. T., Jr., R. J. Smith, Jr., and R. E. Talbert. increased as weed density increased.
1988a. Broadleaf signalgrass (Brachiaria platyphyl- Cramer and Burnside (1982) showed that
la) duration of interference in rice (Oryza sativa). sorghum yield was reduced 4 to 29 percent with
Weed Sci. 36:747750. 11,000 to 45,200 common milkweed plants ha-1.
. 1988b. Interspecific and intraspecific interfer- Common milkweed has a greater effect on sorghum
ence of broadleaf signalgrass (Brachiaria platy- than on corn or soybean yield. Hemp dogbane,
phylla) in rice (Oryza sativa). Weed Sci. another perennial weed, reduced sorghum yield 37
36:589593. to 41 percent, a greater effect than was demonstrat-
Ni, H., K. Moody, R. P. Robles, E. C. Paller, Jr., and ed in corn and soybeans (Schultz and Burnside
J. S. Lales. 2000. Oryza sativa plant traits confer- 1979).
ring competitive ability against weeds. Weed Sci. Weerakoon and Lovett (1986) also found that
sorghum has limited competitive ability in work
Pantone, D. J., and J. B. Baker. 1991. Reciprocal yield
with lanceleaf sage in Australia. The weed was more
analysis of red rice (Oryza sativa) competition in
competitive with a summer crop of sorghum than in
cultivated rice. Weed Sci. 39:4247.
winter wheat.
Pantone, D. J., J. B. Baker, and P. W. Jordan. 1992.
Path analysis of red rice (Oryza sativa L.) Competi- Redroot pigweed at densities of 0.5, 1, 2, 4, or 12
tion with cultivated rice. Weed Sci. 40:313319. plants m-1 of row in a 25-cm band over the crop row
Perera, K. K., P. G. Ayres, and H. P. M. Gunasena. was planted with sorghum or when sorghum was in
1992. Root growth and the relative importance of the 3- to 4-leaf stage (Knezevic et al. 1997). The rec-
root and shoot competition in interactions between tangular hyperbola model, based solely on weed
rice (Oryza sativa) and Echinochloa crus-galli. density, was not well suited to estimate sorghum
Weed Res. 32:6776. yield across locations. A quadratic polynomial equa-
Ransom, J. K., and E. A. Oelke. 1982. Water plantain tion, because it was able to account for time of weed
(Alisman triviale) interference with wild rice (Ziza- emergence relative to crop growth stage, was more
nia palustris). Weed Sci. 30:1014. appropriate. Knezevic et al. (1997) determined that
The Effect of Weed Density 57

the time of weed emergence relative to sorghums cent of the worlds soybeans, about one-third of
leaf growth stage was critical to estimating the which are exported. In 1925 the total U.S. crop was
weeds effect on final sorghum yield. Significant about 5 million bushels. It grew rapidly to 90 million
sorghum yield losses occurred only when redroot bushels in 1939, almost 300 million in 1950, 700
pigweed emerged before sorghum had 5.5 leaves. million in 1963, and more than a billion bushels in
In an experiment with implications relative to cli- 1980. In 1994, U.S. farmers grew 2.5 billion bushels
mate change, weed competition, and crop growth, (United Nations 2000).
Ziska (2001) showed in climate-controlled green- It is also a crop that demands careful weed con-
house studies that single-leaf photosynthetic rates trol. Herbicides are used for weed management in
declined for sorghum and common cocklebur in most fields where soybeans are grown. Total use in
competition. Elevated CO2 reduced the percentage 1997 was 84.5 million pounds or 13.3 percent of the
decline of common cocklebur and increased it in U.S. herbicide market. Sixty-eight percent of the
sorghum 35 days after planting, relative to ambient U.S. crop acreage was planted with genetically
CO2 level. When both plants were grown in mono- modified seed in 2001 (
culture, elevated CO2 significantly stimulated leaf statuniv/att), all of which received at least one her-
photosynthetic rate, leaf area, and aboveground dry bicide application.
weight of common cocklebur more than that of The number of studies of weed-crop interference
sorghum. Therefore, Ziska (2001) concluded that as in soybeans exceeds those for any other crop by at
atmospheric CO2 continues to increase, vegetative least a factor of 2. Stoller et al. (1987) summarized
growth, competition, and potential yield of several the extant work on soybean-weed interference in a
economically important C4 crops could be reduced complete review that is a good starting point for
when they compete with C3 weeds such as common those who wish to study soybean-weed interference.
cocklebur. The majority of studies reports the effect of
known densities of a specific weed on the yield of
Literature Cited
soybeans. Nearly 30 different weeds have been stud-
Cramer, G. L., and O. C. Burnside. 1982. Distribution ied but the most work has been done on common
and interference of common milkweed (Asclepias cocklebur, sicklepod, velvetleaf, and pitted morn-
syriaca) in Nebraska. Weed Sci. 30:385388. ingglory. A few studies have emphasized other fac-
Knezevic, S. Z., M. J. Horak, and R. L. Vanderlip. tors in the interference equation, and these will be
1997. Relative time of redroot pigweed (Amaran- reviewed before turning to those that deal with the
thus retroflexus L.) emergence is critical in effects of specific weeds.
pigweed-sorghum [Sorghum bicolor (L.) Moench] Stoller et al. (1987) reviewed a few papers that
competition. Weed Sci. 45:502508. dealt with differences in the competitive ability of
Schultz, M. E., and O. C. Burnside. 1979. Distribu- soybean cultivars, including early work by Burnside
tion, competition and phenology of hemp dogbane
(1979) who showed that Amsoy 71 was more com-
(Apocynum cannabinum) in Nebraska. Weed Sci.
petitive than Beeson with early- and late-emerging
weeds. Van Acker et al. (1993) suggested, as others
Smith, B. S., D. S. Murray, J. D. Green, W. M.
Wanyahaya, and D. L. Weeks. 1990. Interference of
have, that development of cultivars with early
three annual grasses with grain sorghum (Sorghum branching and the use of narrower rows would be
bicolor). Weed Technol. 4:245249. successful weed management techniques. Shaw et
Weerakoon, W. L., and J. V. Lovett. 1986. Studies of al. (1997) showed that one (Hutcheson) of three
Salvia reflexa Hornem. V. Competition from crop Group V soybean cultivars was consistently more
and pasture species. Weed Res. 26:283290. competitive in Mississippi. Buehring et al. (2002),
Ziska, L. H. 2001. Changes in competitive ability also in Mississippi, found that the same cultivar
between a C4 crop and a C3 weed with elevated car- (Hutcheson) was the only one of three tested that
bon dioxide. Weed Sci. 49:622627. contributed to sicklepod control and increased soy-
bean yield but only in narrow (19 cm) as opposed to
38 cm rows. However, it was only under optimum
Soybeans, one of the worlds major crops, are very growing conditions that narrow rows improved soy-
important to U.S. agriculture. The soybean, native to bean yield. A glyphosate-resistant cultivar grown in
China, is a crop that has grown to prominence in my medium population (455,735 plants ha-1 ) and nar-
lifetime. The United States grows more than 50 per- row (19 cm) rows with two glyphosate applications
58 Chapter 5

gave similar sicklepod control and a 24 percent cultivar had early interference effects on cocklebur
greater yield than soybeans grown in 76 cm rows. that were 5.3 and 9.5 times greater than the other
Shilling et al. (1995) demonstrated, in field stud- indeterminate cultivar, which had a greater interfer-
ies, that sicklepod was tallest when grown with the ence effect in the late period.
tallest cultivars (Centennial or Biloxi) and shortest It is also well accepted that narrow rows increase
when grown with a dwarf isoline of the cultivar soybeans competitive ability, reduce weed compe-
Tracy M. Depending on the cultivar, soybean com- tition, and may increase yield. Costa et al. (1980)
petition reduced early-season sicklepod density 30 found that 27 cm rows versus more conventional
to 50 percent. Centennial, a tall cultivar from matu- (in 1980) 76 cm rows, produced an average seed
rity Group VI, and Tracy M, a short cultivar from yield 21 percent higher over all years, populations,
maturity Group VI, reduced early-season sicklepod and cultivars. The review by Stoller et al. (1987)
biomass 30 percent. Sharkey and Biloxi (maturity stated that when soybeans emerged before vel-
Group VII) reduced sicklepod biomass 40 percent. vetleaf, competition was always reduced to soy-
Late in the season, sicklepod biomass reduction beans benefit. Early planting of soybeans favored
ranged from 18 percent for Tracy M to 55 percent soybeans over velvetleaf primarily due to vel-
for Biloxi and was directly related to cultivar height. vetleafs photoperiod sensitivity. Stoller et al.
It is well accepted that soybean cultivars differ in (1987) citing Murphy and Gosset (1981) also
their competitive ability. Exactly why one is more claimed that planting date did not affect the period
competitive than another remains unknown. Accord- of weed-free maintenance required early in the sea-
ing to Rose et al. (1984), the factors that determine son to prevent yield loss. Parker et al. (1981) found
a soybean cultivars competitive ability (and one little yield response to changes in row width when
assumes the competitive ability of most crop plants) soybeans were planted on time. Planting later than
include: rate of emergence (compared to competing June lowered yield especially in wider rows. In
weeds), seedling vigor (growth rate), rapidity of contrast, Klingaman and Oliver (1994a) found that
canopy closure, and allelopathic attributes. the percent soybean yield loss increased as plant-
James et al. (1988) evaluated 12 soybean cultivars ing date was delayed after early May. Soybean
in competition with sicklepod and found no correla- yield losses from 1.7 weeds m-1 of row were 10, 18,
tion of soybean cultivar maturity group or date of and 20 percent for early-May, mid-May, and early-
introduction as a commonly used cultivar with sick- June plantings. Yelverton and Coble (1991) also
lepod interference. James et al. (1988) recommend- showed that as row spacing increased weed densi-
ed development of cultivars with improved tolerance ty increased and the density and effect of weeds
to sicklepod infestation. Monks and Oliver (1988) coincided closely with the amount of light that
did not find any cultivar advantage in competition penetrated to the soil surface.
with several common southern weeds. Shaw et al. (1991) found that a series of herbicide
Bussan et al. (1997), on the other hand, found that treatments all controlled sicklepod better late in the
the yield and ranking of 16 soybean cultivars varied growing season when soybeans were planted in 25 cm
with the weed with which they competed. Grass as opposed to 97 cm rows. Bendixen (1988) con-
weeds reduced yield the most and small-seeded firmed that better johnsongrass control was achieved
broadleaved weeds reduced yield the least. Some with four of six herbicides when they were used in
cultivars yielded well and still allowed a high weed 25 cm as opposed to 76 cm rows. Walker et al. (1984)
biomass. In their work (Bussan et al. 1997), there found no effect of row spacing on soybean height or
was no relationship between weed competitiveness seed size, but the number of pods per plant was high-
and soybean canopy area, height, and plant volume er in 80 than in 40 cm rows. Soybeans planted in 20
30 to 45 days after planting. Jordan (1992) used and cm rows outyielded those 40 cm and 80 cm rows if
recommended path analysis to study differential sicklepod was not controlled. The work of Walker et
interference between cultivars and weeds. He used a al. (1984) was similar to that of McWhorter and Sci-
semidwarf determinate and an indeterminate culti- umbato (1988), who showed that the height and
var. When the two cultivars grew with common weight of sicklepod were less when soybeans were
cocklebur, there was an early period in interference, planted in 25 cm as opposed to 102 cm rows. Sickle-
40 to 62 days after planting, and a later period, 63 to pod interference for the entire season reduced soy-
145 days after planting. Late interference measures bean yield regardless of the soybean row spacing and
were independent of early measures. The semidwarf the average soybean yield increased with 51 cm rows
The Effect of Weed Density 59

compared to 102 cm rows regardless of the length of the underlying physiology or mechanism is not
sicklepod competition. explored as it should be.
Jackson et al. (1985) found that the time of weed Scott and Geddes (1979) showed that, on a given
removal from soybeans was as important as the extent day, the differences in water potential between soy-
of removal. Interference up to 4 weeks after soybean beans and common cocklebur were small. They
emergence did not reduce soybean yield as long as found greater diffusive resistance values in soybeans
moisture was adequate. With a drought or very high and the diffusive resistance was always greater when
weed density, yield was reduced with 4 weeks of inter- either species was in competition. Patterson (1986)
ference and there was no difference in the competitive showed that growth reduction related to water stress
effect of annual grasses and annual broadleaved was greater for soybeans than for sicklepod. With
weeds. Drought also played a role in work by adequate water, competition from sicklepod reduced
Mortensen and Coble (1989). Well-watered and leaf area duration (LAD) of soybeans while compe-
drought-stressed common cocklebur reduced soybean tition from soybeans reduced LAD and sicklepods
yield 29 and 12 percent, respectively. Drought- net assimilation rate (NAR). In competition studies,
stressed common cocklebur interfered with soybeans sicklepod reduced soybean dry weight more under
over a shorter distance, and the magnitude of the effect drought than with adequate water. In greenhouse
at any distance was reduced. Drought (water stress) studies, Patterson and Flint (1983) found net photo-
was more harmful to common cocklebur than to soy- synthetic rate, net assimilation rate, and water use
beans. Common cockleburs canopy diameter, stem efficiency on a whole plant or single leaf basis were
diameter, node number, and height were all reduced as greatest in the C4 plant smooth pigweed than in soy-
they were in soybeans but the magnitude of the effect beans, common cocklebur, jimsonweed, prickly
was greater in common cocklebur. Soybeans yield sida, spurred anoda, or velvetleaf. Total dry matter
potential was reduced by drought stress and that production 29 days after planting under similar con-
reduction also reduced the effect of weed interference. ditions was greatest in common cocklebur and least
For example, in well-watered soybeans, the canopy in jimsonweed. Munger et al. (1987) found that at
closed about 12 weeks after emergence. In drought- leaf water potentials less than -2.5 MPa, stomatal
stressed soybeans, the canopy never closed and this conductance, net photosynthetic rate, and transpira-
reduced light interference between soybeans and com- tion rate were always greater in velvetleaf than in
mon cocklebur. Mortensen and Coble (1989) conclud- soybeans. Velvetleafs photosynthetic rate increased
ed that reciprocal interference between soybeans and linearly up to 1.5 cm s-1, but there was no further
common cocklebur is not stable across soil-moisture increase above 1.5. As water stress became greater,
conditions, and that was significant for modeling of stomatal conductance, photosynthetic rate, and tran-
the interactions. spiration of velvetleaf declined more rapidly than
Patterson et al. (1988) studied the effect of small they did in soybeans (Munger et al. 1987). Thus, one
(2.7 m2) versus large (11 m2) plots on results with row might conclude that with poor moisture, soybeans
spacings of 15, 30, 45, and 90 cm when soybeans should be a more-effective competitor with late-
competed with sicklepod or common cocklebur. The emerging velvetleaf.
biomass and seed yield of both weeds in small and In a study of the role of water in interference
large plots increased as soybean row spacing in- between common cocklebur, entireleaf morning-
creased. Soybean biomass was not affected by row glory, and soybeans, Mosier and Oliver (1995b)
spacing if weeds were absent. Patterson et al. (1988) showed that interference from soybeans and entire-
suggested that soybean biomass from small plots may leaf morningglory or from soybeans alone reduced
be substituted for seed yield from large plots as a the leaf area index and growth rate of common cock-
measure of sicklepod or common cocklebur interfer- lebur more than entireleaf morningglory alone, and
ence, if both plots have the same row spacing. the effects were always greater without irrigation.
A few reports have emphasized the effect of envi- Irrigated cocklebur produced 687 burs per plant
ronment (e.g., water, light, and temperature) on whereas nonirrigated plants produced only 359. Irri-
interference in soybeans. No reports on nutrient gated entireleaf morningglory reduced cocklebur
competition in soybeans were found. Too often the bur production 42 percent, but without irrigation bur
differences between years in repeated studies are production was reduced only 28 percent. A mixture
said to be due to environmental differences and, in of soybeans and common cocklebur reduced entire-
fact, they are. But the explanation stops there and leaf morningglory seed production 84 to 90 percent.
60 Chapter 5

Entireleaf morningglory is an important weed in bania growth was more stimulated by warmer tem-
soybeans in the southern United States, but it was peratures than soybean growth. Stoller and Myers
not competitive in any treatment combination either (1989b) emphasized the importance of light compe-
year of this study (Mosier and Oliver 1995b) tition in their study of interference between soy-
because of its low leaf area index and, therefore, its beans and four weeds. All five species adjusted to
inability to compete effectively for light and water. reduced irradiance by decreasing the rate of light-
In a separate report, Mosier and Oliver (1995a) saturated photosynthesis, photosynthetic leaf respi-
showed that both weeds were effective competitors ration rates, root shoot ratios, and leaf density while
in soybean. The total leaf area index, the leaf area increasing their leaf area ratio. As irradiance was
index within the soybean canopy, and soybeans reduced, plant support tissues (roots, stems, and
growth rate and seed yield were decreased more by petioles) and leaf ratios did not change for common
common cocklebur than by entireleaf morningglory. lambsquarters or velvetleaf. All increased for soy-
The essential role of water in interference is illus- beans, eastern black nightshade, and tumble pig-
trated by data that showed that interference from weed. This indicates a superior adaptation of the
entireleaf morningglory, common cocklebur, or both latter two weeds for efficient light harvesting with
reduced soybean yield 21, 57, or 64 percent with reduced light. Of the five species studied, eastern
irrigation and 12, 60, or 76 percent without irriga- black nightshade had the lowest respiration rate, the
tion. Soybeans extracted water from greater soil highest leaf area ratio, and the lowest support tissue
depths when it grew with weeds than when it grew to leaf area ratio and was optimally adapted for
in monoculture. Mosier and Oliver (1995a) conclud- superior competition under reduced light (i.e., under
ed that soybeans high water use efficiency (WUE) the soybean canopy).
without irrigation suggests that soybeans use water When the lower leaves of greenhouse-grown
more efficiently when soil water is limiting than common cocklebur and velvetleaf were shaded to
when it is abundant. only 5 percent of full light for 12 days, there was an
Geddes et al. (1979) explained some of the com- increase in upper (unshaded) leaf area beginning 3
petitive effects of cocklebur for water. Roots of (velvetleaf) and 6 (common cocklebur) days after
common cocklebur explored a greater volume of soil shading began (Regnier and Harrison 1993). Total
than did those of soybeans. The total amount of water plant dry weight of velvetleaf 12 days after shading
used by a pure stand of soybeans and a mixed stand began was unaffected by shading, but it was reduced
of soybeans and common cocklebur was greater than 10 percent in common cocklebur. Regnier and Har-
that used by a pure stand of common cocklebur. rison (1993) showed that common cocklebur has
Twelve weeks after emergence, the percent reduction greater shade tolerance than velvetleaf and that both
in dry matter and leaf area due to interspecific com- species have the ability to compensate for shading of
petition were greater in common cocklebur than soy- lower leaves by altering upper shoot growth.
beans and they were greater in a dry than in a wet Murphy and Gosset (1981) found shading
year. There was no difference in water use efficiency increased until 11 weeks after planting and then
(WUE) between the species in a wet year and for up declined (there was more light at the soil surface) 14
to 10 weeks after emergence in a dry year (Geddes et weeks after planting because of soybean leaf loss.
al. 1979). Light at the soil surface, 3 and 5 weeks after plant-
Cool early season temperature slowed growth of ing averaged 55 and 40 percent of available light,
hemp sesbania but did not affect soybean growth respectively. Murphy and Gosset (1981) found that
and yield. There was negligible competition less shading was required to prevent weed establish-
between soybeans and hemp sesbania densities of 3 ment than the 90 percent previously reported by
or 6 plants m-2 (King and Purcell 1997). In one year, Knake (1972) for control of giant foxtail.
hemp sesbania grew above the soybean canopy and Flint and Patterson (1983) studied temperatures
decreased soybeans light interception 29 to 68 per- effects on growth of soybeans, common cocklebur,
cent and eventual yield 30 to 48 percent. King and and smooth pigweed. For all three species, height,
Purcell (1997) concluded that competition for light dry weight, and leaf area increased significantly
was the primary cause of soybean yield loss from when temperatures ranged from 26/17, 29/20, to
hemp sesbania competition. Both plants increased 32/23C. The net assimilation rate of all three
their dry weight when day/night temperatures were species peaked when temperatures were 29/20C.
30 and 20C compared to 25 and 15C. Hemp ses- Dry weight and leaf area were reduced by interfer-
The Effect of Weed Density 61

ence. Rising temperature reduced the effects of almost always act as stressors on the crop while they
interference on growth of smooth pigweed (a C4 relieve stress from weed competition. One labeled
plant) but not of the other species. Common cockle- herbicide combination reduced soybean height 6
bur and soybeans were roughly equal competitors at percent 21 days after application but did not increase
all temperatures, and both were superior to smooth time to maturity, population density, or yield.
pigweed, which competed less well in any mixture Banks et al. (1986) demonstrated that soybean
at any temperature, but especially at low tempera- yields were similar in no-till and conventionally
tures (Flint and Patterson 1983). Thus, Flint and Pat- tilled plots if sicklepod was absent. If sicklepod was
terson (1983) concluded that common cocklebur is present, soybean yields were higher in no-till plots
more likely than smooth pigweed to compete effec- and increasing sicklepod density caused lower soil
tively in early-seeded soybeans, especially if they water content in tilled and no-till plots.
emerge together. Norsworthy and Oliver (2002a) demonstrated that
One report (Black et al. 1996) showed an effect of glyphosate increased soil moisture availability for
soybean infestation by Rhizoctonia solani in one dryland soybeans and as soybean population
year. Independent of infestation by common cockle- increased from 247,000 to 729,000 plants ha-1 pitted
bur, hemp sesbania, or johnsongrass, soybean yields morningglory and hemp sesbania control increased
decreased up to 18 percent. There was no significant from 60 to 91 percent. Three sequential glyphosate
interaction between R. solani infestation and weed applications reduced pitted morningglory seed pro-
density in either year of the study. duction from 247,000 ha-1 to 9,000 and eliminated
A few studies have combined the effects of herbi- hemp sesbania seed production. The work (Nors-
cides with other aspects of competition. Adcock et worthy and Oliver 2002a) illustrates the importance
al. (1990) reported that increasing herbicide rates of consideration of crop planting density and soil
resulted in higher soybean:weed fresh weight ratios moisture in any management plan.
and higher herbicide response coefficients. The spe- If the number of papers dealing with a weed is a
cific ratio was affected by the weed and herbicide good indicator of the weeds importance, then sick-
combination, which is to say that the effect was not lepod and common cocklebur have to be the most
constant over all herbicides or herbicide combina- important weeds in soybeans in the last 20 or so
tions. Mulugeta and Boerboom (2000) reported that years.
weed control efficacy and crop yield were influ- Soybeans that were kept free of sicklepod for 4
enced more by glyphosate application time than by weeks after emergence produced yields equal to
the rate applied. The critical time of weed removal, season-long control over 3 years (Walker et al. 1984;
the time beyond which weed competition reduced Rushing and Oliver 1998). In contrast, McWhorter
soybean yield more than 3 percent compared to a and Sciumbato (1988) found that sicklepod compe-
weed-free check, was at the low rate (0.42 kg ha-1 ) tition for 4 weeks decreased soybean yield in 2 of 3
and soybeans V2 growth stage with 18 cm rows in years. A reason for sicklepods competitiveness in
reduced tillage plots and at V2 in 76 cm rows and soybeans is that whereas soybeans grew only slight-
with reduced tillage for both years of the study. The ly between 35 and 84 days after emergence, sickle-
critical time of weed removal (with glyphosate in pod continued to grow until it was 30 to 45 cm taller
this study) for 18 cm and 76 cm rows in no-tillage than soybeans (Bozsa et al. 1989). This is explained
soybeans was at the V4 stage of soybean growth. at least partially by the fact that sicklepods growth
The study (Mulugeta and Boerboom 2000) showed responded to distance from the soybean row and
that the critical time of weed removal varied time of emergence relative to the crop (Smith and
between reduced- and no-tillage and between years. Jordan 1993). In a separate greenhouse study, Bozsa
A single glyphosate application can prevent yield and Oliver (1990) showed that soybeans were 1.5 to
loss in narrow-row glyphosate-resistant soybeans 2 times taller than common cocklebur and were
when growing conditions are favorable, but applica- more competitive aboveground during the first few
tion timing is more critical with wide rows because weeks of growth. Common cocklebur had 20 to 50
the critical period for weed removal occurs early in percent more small roots, which have a greater
the growing season. A second herbicide application uptake of water and nutrients per unit of surface area
may be needed for later-emerging weeds in widely than larger roots, and it was more competitive below
spaced soybeans. Krausz et al. (2001) demonstrated ground during early growth (Bozsa and Oliver
what is often assumed but rarely shown: herbicides 1990). Thus, by 4 weeks after emergence, common
62 Chapter 5

cockleburs greater capacity for root competition However, these results contrast with those of Berti
significantly reduced soybeans growth. and Sattin (1996) who found that the position of
These observations are at least partially explained common cocklebur or barnyardgrass relative to the
by the fact that common cocklebur has a longer veg- soybean row was of little importance relative to
etative growth period than soybeans and was twice weed density as a determinant of yield loss. Relative
as tall at maturity (Bozsa and Oliver 1993). Shoot cover was the most important factor. Both common
and root dry weight and seed yield of soybeans were cocklebur and barnyardgrass can grow taller than
reduced by whole-plant (shoot and root) interfer- soybeans. Berti and Sattin (1996) concluded that for
ence from common cocklebur, but common cockle- weeds that grow taller than the crop, the main com-
bur growth was not affected by soybeans presence. petitive factor is shading caused by the taller plant.
Only common cocklebur shoot interference reduces Interference of one common cocklebur in 1.8, 0.9,
soybean seed production 48 percent, which was or 0.3 m of row for 8 weeks reduced soybean yield
equal to that caused by whole-plant interference 7, 14, or 30 percent and full-season interference
(Bozsa and Oliver 1993). Regnier et al. (1989) had reduced yield 16, 33, or 65 percent (Rushing and
shown previously that decreases in soybean yield Oliver 1998). If one common cocklebur 3 m-1 of
were due to shoot interference. Yield always crop row interfered for the whole season, soybean
decreased more when root interference also yield was reduced 3 to 12 percent (Bloomberg et al.
occurred, but the shoot effects and competition for 1982). Soybean yield, total dry weight, and pods per
light dominated as a cause of common cockleburs plant all increased as the length of time between
effects on soybeans. soybeans and common cocklebur emergence
Regnier and Stoller (1989) demonstrated that increased. If common cocklebur emerged 4 weeks
common cocklebur had more leaves within the soy- after soybeans, yield was reduced 7 percent. When
bean canopy than jimsonweed or velvetleaf. By the common cocklebur was removed 6 weeks after soy-
end of the growing season, common cockleburs leaf bean emergence, soybean yield was reduced less
area was evenly distributed above and below the than 10 percent (Bloomberg et al. 1982).
soybean canopy while nearly all the leaf area of the Monks and Oliver (1988) studied the interaction
other two weeds was above the soybean canopy. between two soybean cultivars and common cockle-
Regnier and Stoller (1989) proposed that common bur, johnsongrass, Palmer amaranth, sicklepod, and
cocklebur had greater shade tolerance than the other tall morningglory. There was no reduction in soy-
two weeds. Growth from the lower axillary buds of bean biomass for 6 weeks after emergence. Only
jimsonweed and velvetleaf was strongly inhibited by common cocklebur and Palmer amaranth reduced
soybeans but was not for common cocklebur. The soybean biomass during the growing season. The
latter had more axillary growth along lower stems other three weeds grew more slowly and had no
than soybeans, which makes it a more aggressive measurable effect on soybean biomass. Soybean,
competitor even though common cocklebur and soy- however, was an effective competitor because it
beans were similar in height and seemed to compete reduced the biomass of all weeds 90 to 97 percent.
for the same aboveground niche. Lower branching The biomass of both soybean cultivars was reduced
and shade tolerance make common cocklebur a when they grew within 50 cm of Palmer amaranth.
more effective competitor (Regnier and Stoller Soybean seed yield as distinct from biomass was
1989). reduced when the soybean grew within 25 cm of
Sicklepods height, the number of main stem common cocklebur and Palmer amaranth or within
nodes, the number of branches, and its shoot dry 12.5 cm of the less-competitive tall morningglory
weight all decreased 12 weeks after emergence (Monks and Oliver 1988). Study of competition of
when the plants were close to the soybean row Palmer amaranth alone showed soybean yield was
(Bozsa et al. 1989). Plants that emerged 7 days after highly correlated with Palmer amaranth biomass 8
soybeans were shorter. Nearly all sicklepod plants weeks after soybean emergence and to the weeds
were taller than soybeans, but if they were close to density (Klingaman and Oliver 1994b). Palmer ama-
the soybean row, their dry weight was reduced up to ranth densities of 0.33, 0.66, 1, 2, 3.33, and 10
60 percent. Similarly, sicklepods that were 10 to 30 plants m-1 of row reduced soybean yield 17, 27, 32,
cm apart in the soybean row reduced soybean yield 48, 64, and 68 percent, respectively. Soybean yield
25 to 35 percent more than when they were 90 cm reduction was approximately linear to about 2
apart (Bozsa et al. 1989). Palmer amaranth m-1 of row suggesting that
The Effect of Weed Density 63

intraspecific interference between adjacent weeds delayed harvest, but that advantage was offset when
began at relatively low weed densities. johnsongrass was present. Hemp sesbania reduced
The interaction between insect defoliation and soybean yield of the early maturing cultivar 23 per-
weed interference was studied by Grymes et al. cent when soybean was harvested 1 week after matu-
(1999). Interference was with common cocklebur, rity and 26 percent when it was harvested on the other
hemp sesbania, or johnsongrass at 2.5, 0.5, or 2 two dates. The late-maturing cultivars yield was
plants m-2, respectively. Defoliation to simulate in- reduced 16, 22, or 28 percent when it was harvested
sect action, at R2 (full bloom) and R5 (beginning 1, 2, or 3 weeks after maturity. Late harvest tended to
seed development), was done by removing 1 or 2 decrease the value of the soybean yield because of
leaflets from each soybean trifoliate leaf, which increased foreign material, increased moisture, and
approximated 33 or 66 percent defoliation. The damaged kernels (McWhorter and Anderson 1993).
weeds were not affected by this. Soybean height While johnsongrass has not received as much
3 weeks after defoliation at R5 was not influenced by research attention as common cocklebur, it has been
weed interference, soybean defoliation level, or and remains an important weed in soybeans. Vitta
defoliation stage. Averaged across defoliation levels and Satorre (1999) showed why johnsongrass is usu-
and growth stages, johnsongrass, common cockle- ally more competitive early in the growing season
bur, and hemp sesbania reduced soybean yield 30 to by evaluating canopy characteristics. They found a
35, 15, and 14 percent, respectively. As soybean significant linear relationship between the relative
defoliation level increased, there was a linear leaf area of johnsongrass and its contribution to the
decrease in soybean yield. For all three weeds and total biomass of the mixture of soybeans and john-
both defoliation stages, 33 and 66 percent defolia- songrass, measured early in the growing season. In
tion reduced soybean yield 6 and 20 percent in one monoculture, crop and weed canopies developed
year and 12 and 33 percent in the second year. Defo- simultaneously. They began to compete at a thermal
liation at R5 reduced yield 10 percent more than time of 250 to 350C days after sowing, which cor-
defoliation at R2 in one year (Grymes et al. 1999). responded with the beginning of the active net
Sims and Oliver (1990) compared interference growth period of johnsongrass biomass. Vitta and
from johnsongrass and sicklepod with and without Satorre (1999) claimed that the relationship between
irrigation. Johnsongrass reduced soybean growth soybean yield and the weeds leaf area was always
early in the season, whereas sicklepod was the more linear but that the slope varied with crop sowing
effective competitor and was competitive all season date. Their results suggest that measurement of
with the effect being greatest during soybeans johnsongrass leaf area may be a simple, effective
reproductive stage. Soybean seed yield was reduced way of predicting soybean yield loss.
31 percent by full-season interference from sickle- Johnsongrass was a less-effective competitor than
pod, 14 percent by johnsongrass, and the combina- smooth pigweed because the latter captured 1.8 to
tion reduced yield 36 percent. Johnsongrasss dry 2.5 times more light and produced more dry matter
matter and seed yield were reduced more than that at all densities of multispecies populations of
of sicklepod by soybean interference. Soybean plus smooth pigweed, johnsongrass, and soybeans (Toler
sicklepod reduced johnsongrass seed production 73 et al. 1996). Nevertheless, johnsongrass presence is
to 95 percent. Sicklepod produced 6 to 31 percent not to be neglected. The number of johnsongrass
fewer seeds when it grew with johnsongrass and 47 culms at harvest was more correlated with soybean
to 75 percent fewer seeds when it grew with soy- yield loss than the number of plants per unit area 4
beans or soybeans and johnsongrass (Sims and Oliv- to 6 weeks after planting (Williams and Hayes
er 1990). 1984). Full-season johnsongrass competition
McWhorter and Anderson (1993) demonstrated reduced soybean yield 59 to 88 percent, and soy-
that johnsongrass was slightly more competitive than beans could not tolerate what Williams and Hayes
hemp sesbania with an early-maturing soybean culti- (1984) called a heavy infestation for more than 5
var. Johnsongrass reduced soybean yield 32, 35, and weeks after planting without a yield loss.
36 percent over 3 years when soybeans were harvest- Interference of velvetleaf with soybeans has also
ed 1, 2, or 3 weeks after maturity. A late-maturing been studied frequently. Higgins et al. (1984) found
cultivars yield was reduced 27, 29, or 39 percent that monocultural velvetleaf consistently exceeded
when it was harvested 1, 2, or 3 weeks after maturity. velvetleaf grown with soybeans in leaf area, nodes
The seed grade of both cultivars improved with with fully developed leaves, canopy width, branches,
64 Chapter 5

and number of capsules as early as 3 weeks after which seems counterintuitive. This was related to
emergence. Velvetleaf without soybean competition velvetleafs increasing interception of light with
developed more than nine times the dry matter of vel- increasing height. Reductions in soybean yield and
vetleaf grown with soybean. When soybeans were par- yield components was greater if velvetleaf was
tially defoliated to simulate damage from the green maintained at various heights for only 3 weeks
cloverworm (Plathypena scabra F.), the leaf area, rather than 6 weeks following soybean emergence.
number of leaves, and number of main stem nodes of This was attributed to a longer duration of light
velvetleaf decreased. Because the green cloverworm competition by velvetleaf. Even when the weeds
attacks soybeans late in the growing season, velvetleaf height was kept 25 percent below the soybean
will obtain only a slight advantage and will have done canopy for 4 weeks, soybean yield and pod and
its damage before the green cloverworm appears. In branch numbers decreased when velvetleaf was
spite of the fact that soybeans compete well with vel- allowed to regrow for the rest of the season, but this
vetleaf, the latter is regarded as a major problem in was not true if the weed was removed after the first
most U.S. soybean-producing areas. 4 weeks. Soybean yield was not affected if vel-
Velvetleaf is taller than soybeans for most of the vetleaf plants that were 25 percent shorter than soy-
growing season and has more branches, especially beans were removed or allowed to regrow after an
near the top of the canopy (Akey et al. 1990). Vel- initial 6 to 8 weeks of clipping (Begonia et al. 1991).
vetleaf therefore has greater light interception abili- Dekker and Meggitt (1983a) showed that the
ty than soybean, especially early and late on any effects of low populations (2.4 to 4.7 plants m-2) of
day. The leaf canopies of velvetleaf and soybeans velvetleaf were greatest on soybean yield and less
had similar total light interception on most sample on flowering node and dry weight production. Vel-
days, but velvetleaf had higher light utilization effi- vetleaf exerted its effect by its presence and not by
ciency (Akey et al. 1991). That is, velvetleaf con- changes in its density. Further work (Dekker and
verted more of the intercepted light energy to dry Meggitt 1983b) showed that velvetleaf had the adap-
matter, especially with its emergent canopy in the tive ability of differential mortality at different pop-
middle and late parts of the growing season. To fur- ulation densities and soybean did not. The result is
ther explain velvetleafs competitive ability, Akey et that the velvetleaf plants that remain grow large and
al. (1991) studied the relative competitive ability of are more competitive as the population declines due
velvetleaf and soybeans. The relative aboveground to death. Soybean plants do not die and release
dry weight of soybeans in mixtures was higher than resources for the survivors; they become smaller and
expected from monocultural values early in the sea- less competitive.
son and lower than expected late in the season. Vel- Velvetleafs effects can be mitigated by adjusting
vetleaf, on the other hand, had higher than expected soybeans planting date (Oliver 1979). Velvetleaf
values late in the season. Velvetleaf depressed seed was planted at one plant per 61 or 30 cm of row with
yield of soybeans in all mixtures, and the relative competition ranging from 4 weeks to full season.
seed yield of velvetleaf was greater in all mixtures Velvetleaf that emerged with soybeans in mid-May
than in monoculture. Soybeans relative growth rate was twice as competitive as velvetleaf that emerged
(RGR), leaf area ratio (LAR), and net assimilation with soybeans planted in late June. One velvetleaf
rate (NAR) did not differ significantly among mix- per 30 cm of soybean row competing for the full
tures and decreased over the season. Velvetleaf did season reduced soybean yield 27 percent for the
not show any significant differences in RGR, LAR, early (mid-May) planting and only 14 percent for
or NAR over the growing season. Velvetleafs RGR the late-June planting. Oliver (1979) attributed the
and LAR were highest early in the growing season difference to the short-day photoperiodic response
and progressively declined. Velvetleaf is a good of velvetleaf. Early growth stages of soybeans are
competitor, in part because it has a higher NAR and competitive with velvetleaf. Ten weeks after emer-
RGR early in the season even though competition is gence, velvetleaf competition reduced soybean
not significant then. During that time, velvetleaf growth and development. Oliver (1979) postulated
gained resources at the expense of soybeans that that velvetleaf may not be an important problem in
enabled it to compete effectively in midseason and Arkansas (southern United States) because of its
late season. photoperiodic response and late-season competitive-
Begonia et al. (1991) postulated that soybean ness. However, when soybeans are planted early to
yield was inversely related to velvetleafs height, gain a yield advantage, velvetleaf has proven to be
The Effect of Weed Density 65

an important weed. In slight contrast to several Common LambsquarterChenopodium album

papers reported above, Munger et al. (1987) sug-
With 32 common lambsquarters 10 m-1, 10 weeks of
gested that interspecific competition for soil water
interference were required to cause a 20 percent
played an important role in the interactions between
yield reduction when weeds were removed by hand.
velvetleaf and soybeans but that the competitive
If a postemergence herbicide was used to kill the
interactions were due to resource limitations other
weed, a 20 percent yield reduction occurred if it was
than water. They showed that monocultural vel-
not applied prior to 5 weeks after emergence (Crook
vetleaf with 5 plants m-2 extracted water up to 1 m
and Renner 1990). The authors attributed the differ-
deep, whereas monocultural soybeans (32.5 plants
ence to the fact that hand removal gave complete
m-2) extracted water up to 1.5 m or more. Interspe-
control whereas the herbicide did not. Work in Ohio
cific competition between soybeans and velvetleaf
(Harrison 1990) showed that common lambsquar-
resulted in a 40 percent reduction in soybean seed
ters was a more vigorous competitor. When 5 per-
yield and a 50 percent reduction in velvetleaf seed
cent yield loss was used as the threshold, regression
yield. Interspecific competition had little to no effect
analysis predicted a threshold density of 2 weeds
on soybean morphology before 8 weeks after plant-
m-1 5 weeks after emergence and 1 weed m-1 7 weeks
ing (Munger et al. 1987).
after emergence. Each kg ha-1 of weed biomass
Once acquired in a field, velvetleaf continues to
resulted in an average soybean yield loss of 0.26 kg
be important because only 6.8 +- 0.5 percent of the
ha-1 (Harrison 1990).
soil seedbank emerges each year (Lindquist et al.
1995). In the absence of crop competition, a vel- Common MilkweedAsclepias syriaca
vetleaf plant could produce 125 to 227 seeds. Vel-
vetleaf that emerged early produced the largest The average yield reduction of soybeans from
number of seeds, but the seed production declined 11,000 to 45,000 common milkweed plants ha-1 was
up to 82 percent with crop competition. 12 to 19 percent, slightly higher than for corn
(Cramer and Burnside 1982).
Other Weeds
Common RagweedAmbrosia artemisiifolia
If the number of papers in print is an appropriate cri-
terion, then the preceding weeds (common cockle- The damage threshold for full-season, in-row inter-
bur, velvetleaf, and johnsongrass) are the most ference was 4 common ragweed 10 m-1 of row and
important weeds in soybeans. The first edition of they caused an 8 percent yield loss (Coble et al.
this book (Zimdahl 1980) noted that common cock- 1981). Soybean yield was not reduced by a natural
lebur was the most important and detrimental weed population of common ragweed if the period of
in soybeans. Johnsongrass and velvetleaf studies interference was 6 weeks or less after emergence.
were also reported in the first edition. The first edi- Soybeans kept weed free for 2 weeks or longer after
tion reported on studies of 15 different weeds; more emergence in a dry year produced normal yields but
(28) are included here. Some were reported in a sin- 4 weeks of weed-free maintenance was required
gle manuscript and others were studied in three or when water was adequate early in the growing sea-
four separate reports. There is no particular pattern son. Eight weeks after emergence, common rag-
of the research or the results, except that all weeds weed averaged 25 cm taller than soybeans, and the
reduce soybean yield. These studies are summarized weed intercepted 24 percent of the incident radia-
below in alphabetical order by the weeds common tion. Coble et al. (1981) used these data to determine
name. the economic threshold (when to control) for com-
mon ragweed in soybeans.
BurcucumberSicyos angulatus
Common SunflowerHelianthus annuus
Burcucumber emergence was greatest in late May
through mid-June and had almost ceased by early Soybeans required 4 to 6 weeks free of common
July (midsummer) regardless of the tillage system sunflower to obtain maximum yield (Irons and
(no-till and reduced tillage) or the row spacing (38 Burnside 1982). In Kansas, soybean yield reduc-
and 76 cm) (Esbenshade et al. 2001). Preplant tions ranged from 17 to 19 percent with 0.3 common
tillage increased burcucumber emergence 70 to 110 sunflowers m-2 and 95 to 97 percent with 4 to 6
percent compared to no-tillage, but row spacing had plants m-2 (Geier et al. 1996). Interference was pri-
no effect on emergence or biomass production. marily for light. Evidence was provided to show that
66 Chapter 5

0.3 common sunflower m-2 reduced photosyntheti- Table 5.2. Effect of Shading on Shoot and
cally active radiation (PAR) at the plant canopy by Seed Production by Eastern Black Nightshade
300 to 390 mol m-2s-1 or 18 to 24 percent. The abil-
Time Light Shoot Berry
ity of common sunflower to intercept PAR above the
(weeks) condition production (g) production (No.)
soybean canopy is, in the authors (Geier et al. 1996)
view, an important component of its interference
20 full sun 243 5957
capability in soybeans. These findings agree with
11 full sun 38 576
those of Allen et al. (2000). They found that soybean
20 94% shade 3 23
yields tended to decrease as the weed-free period
11 94% shade 1 1
occurred later in the growing season and soybeans
had a larger canopy. Early-season weed-free periods Source: Stoller and Myers (1989a).
(2 to 4 and 4 to 6 weeks after planting) allowed
common sunflower to become established before
soybeans had a well-developed canopy. Yield reduc- (table 5.2). Eastern black nightshade plants that
tions were 15 to 80 percent. If the weed-free period emerged with soybeans and were between 75-cm
occurred 6 to 8 or 8 to 10 weeks after planting, the rows produced 43 g of shoots and 264 berries. Those
effect on yield was minimal because the common that grew in the row and emerged 6 weeks after soy-
sunflowers survived only a few weeks after estab- beans produced only 1 g of shoot and 16 berries.
lishment and they did not produce seed. Further illustration of the importance of light is
shown by their finding that shoot growth and berry
Cutleaf GroundcherryPhysalis angulata
production increased from 80 to 200 percent in 2
Cutleaf groundcherry was identified as an inconse- weeks between the initiation of soybean leaf abscis-
quential and easy to control weed in work by Bell sion and maturity.
and Oliver (1979). At densities as high as 60 weeds
Florida BeggarweedDesmodium tortuosum
m-1 of row, in the row, there were no significant
reductions in soybean leaf area index, height, dry The influence of water stress on interference is well
weight, growth rate, or seed yield. illustrated by the work of Griffen et al. (1989). Soy-
bean leaf area and aboveground biomass were
Eastern Black NightshadeSolanum ptycanthum greater than that of Florida beggarweed under opti-
One study emphasizes the effect of soybeans on mum water conditions in a greenhouse study but
eastern black nightshade seed production (Quaken- were equal to or less than the weeds under water
bush and Andersen 1984). Without soybean interfer- stress. Soybeans were more competitive with ade-
ence, one eastern black nightshade planted in May quate soil moisture but less competitive than Florida
produced 7,000 berries and 800,000 seeds. If a plant beggarweed under water stress. In short, water stress
was planted alone in mid-July, it produced up to 100 favored the weed over soybeans.
berries and if planted in August it produced no Giant RagweedAmbrosia trifida
berries. However, when eastern black nightshade
was planted with soybeans in May, a plant produced The injury threshold for giant ragweed was less than 2
less than 85 berries. If planted in June, berry pro- per 9 m-1 of row. Full-season interference at this densi-
duction dropped to none to 3 and a July planting ty reduced soybean yield 46 to 50 percent, and the crit-
produced no berries. If soybeans were defoliated in ical duration was 2 to 4 weeks in one year and 4 to 6 in
July to simulate hail damage, then an eastern black a second year (Baysinger and Sims 1991). Webster et
nightshade plant that had been planted in May pro- al. (1994) attributed most of giant ragweeds competi-
duced up to 1,600 berries, and those planted in July tive effect to its ability to initiate and maintain axillary
produced up to 58 berries. Quakenbush and Ander- leaves and branches within the soybean canopy. They
sen (1984) concluded that one need only control determined that the economic threshold was 0.03 to
eastern black nightshade through June to prevent 0.08 giant ragweed plants m-2.
berry (and seed) production if there is no subsequent
Green Foxtail and Fall PanicumSetaria viridis
hail damage.
and Panicum dichotomiflorum
A second study emphasized the effects of shading
on interference (Stoller and Myers 1989a). Shade Soybeans that were free of a natural mixed stand of
clearly suppressed eastern black nightshade growth giant green foxtail and fall panicum for 2 weeks
The Effect of Weed Density 67

after soybean emergence yielded the same as plots sity. Glyphosate-treated hemp sesbania did not
that were weed free for the growing season (Harris affect soybean yield. Illustrative of the importance
and Ritter 1987). If the grasses grew with soybeans of crop population shown by so many studies, when
for 8 weeks, soybean yield was reduced because of yield loss was averaged over all soybean popula-
a decrease in pods per plant. With a drought, yield tions, it was reduced from 44 to 22 percent by more
reduction did not occur until the weeds had grown than doubling soybean population from 217,000 to
with soybeans for 12 to 16 weeks after emergence. 521,000 plants ha-1.
Hand-established weed densities of 1 grass plant per
ItchgrassRottboellia cochinchinensis
7.5 cm-1 of soybean row reduced yield 0 to 11 per-
cent. Natural grass infestations of unspecified densi- Lejeune et al. (1994) used area of influence proce-
ty present for the season reduced soybean yield 21 dures to evaluate interference of itchgrass and soy-
to 41 percent (Harris and Ritter 1987). beans over 2 years. Soybean seed weight within 20
cm of itchgrass was reduced 15 to 21 percent. In one
Hemp DogbaneApocynum cannabinum
year, seed weight reduction of 9 percent was detect-
Hemp dogbane densities of 28 to 40 shoots m-2 ed 40 to 60 cm from an itchgrass plant. Weight
reduced predicted soybean yield 58 to 75 percent or reductions were attributed to decreases in seed num-
62 to 94 percent with the rectangular hyperbolic or ber of 12 to 22 percent within 40 cm of a weed com-
linear regression models, respectively (Webster et pared to a control area. Itchgrass interference
al. 2000). The study was done over 3 site years and increased soybean height within 40 cm of a weed,
differences between sites were attributed to rainfall but soybean canopy width was not affected. Soy-
and temperature. There was delayed soybean bean interference did not affect height of itchgrass
canopy closure and higher yield loss when soil plants but reduced stem numbers 89 to 94 percent
moisture remained high and temperatures were rela- compared to weeds growing alone. This finding
tively cool. When the two predictive models were affirms that one of the best ways to reduce weed
applied to field populations of hemp dogbane, infestation is to plant a crop. When itchgrass inter-
between 19 and 36 percent and 20 and 29 percent of ference was 8 weeks or less, itchgrass fresh and dry
soybean yield loss could be expected from within weight were similar when the weed grew alone or in
hemp dogbane patches for the rectangular hyper- the soybean row. Both were reduced 80 percent
bolic and linear regression models, respectively. when itchgrass competed with soybeans for 10
Webster et al. (2000) concluded that while the rec- weeks (Lejeune et al. 1994).
tangular hyperbolic model appeared to describe the
Ivyleaf MorninggloryIpomoea hederacea
relationship between soybean yield loss and hemp
dogbane density accurately, the relationship was Ivyleaf morningglory was grown at 1 plant per 7.5,
dominated by the models initial linear phase and 15, 30, 60, or 90 cm of row in 2 years. Each density
may be inappropriate. Schultz and Burnside (1979) competed for 22 to 46 days after planting or for the
observed high infestations of hemp dogbane in soy- full season in one year and for 29 to 60 days after
beans in Nebraska and yield losses from 28 to 41 planting or the full season in the second year
percent from season-long infestation. (Cordes and Bauman 1984). The best indicators of
the competitive effect were changes in leaf area
Hemp SesbaniaSesbania exaltata
index, dry weight, and yield. Ivyleaf morningglory
Norsworthy and Oliver (2002b) studied interference similar to pitted morningglory exerted its greatest
of hemp sesbania in drill-seeded, glyphosate-resistant competitive effect during soybeans reproductive
soybeans in an experiment nearly identical to their stages. The primary competition was apparently for
work on pitted morningglory (Norsworthy and nutrients because photosynthetic irradiance mea-
Oliver 2002c). Soybean densities were 217,000, surements and soil water measurements showed that
371,000, and 521,000 plants ha-1. Hemp sesbania ivyleaf morningglory did not compete for light or
densities were 0, 4, 10, or 16 plants m-2 with and water. All densities of the weed could compete for
without glyphosate applied at the V4 and V6 soy- 46 days after emergence in one year and 90 days in
bean growth stage. Soybean seed yield was reduced the second year without reducing soybean yield, but
43 percent by full-season interference of 16 untreat- full-season competition from 1 weed m-1 of row
ed hemp sesbania m-2, which was less than the effect reduced yield 13 percent in one year and 36 percent
of pitted morningglory (62 percent) at the same den- in a second year (table 5.3). The weeds effect was
68 Chapter 5

Table 5.3. Effect of Several Weeds on Soybean Yield

Weed species Density Yield reduction Source

Common cocklebur One 1.8 m-1 row 7% Rushing and Oliver 1998
One 0.9 m-1 row 14%
One 0.3 m-1 row 30%
Common cocklebur One 3 m-1 row full-season 3 to 12% Bloomberg et al. 1982
Common sunflower Full season 47 to 72% Allen et al. 2000
With irrigation Mosier and Oliver 1995a
Entire leaf morningglory 21%
Common cocklebur 57%
Both 64%
Without irrigation
Entire leaf morningglory 12%
Common cocklebur 60%
Both 76%
Hemp dogbane Full season 28 to 41% Schultz and Burnside 1979
Hemp sesbania 16 m-2 -full season 43% Norsworthy and Oliver 2002b
Jerusalem artichoke Full season
1 tuber m-1 of row 31% Wyse et al. 1986
2 tubers m-1 59%
4 tubers m-1 71%
4 tubers m-1 for
4 weeks after planting 9% Wyse et al. 1986
6 weeks after planting 10%
8 weeks after planting 38%
20 weeks after planting 82%
Jimsonweed 0.3 m-1 of row, full-season 8% Kirkpatrick et al. 1983
1.6 m-1 of row,
2 weeks 7%
4 weeks 14%
full season 41%
Johnsongrass Full season 59 to 88% Williams and Hayes 1984
Johnsongrass with early 1 week after soybean 32% McWhorter and Anderson
maturing cultivar with early maturity 1993
2 weeks after soybean 35%
3 weeks after soybean 36%
Johnsongrass with a late 1 week after maturity 27% McWhorter and Anderson
maturing cultivar 2 weeks after maturity 29% 1993
3 weeks after maturity 29%
Ivyleaf morningglory 1 plant 15 cm of row 13 to 36% Cordes et al. 1984
full season
Pitted morningglory 1 plant 10 m-2 full season 47% Norsworthy and Oliver 2002c
16 m-2full season 62%
62 m-2full season 81%
Quackgrass Natural stand for Young et al. 1982
6 weeks 11%
8 weeks 23%
Full season 33%
Sicklepod Full season 31% Sims and Oliver 1990
Johnsongrass Full season 14%
Sicklepod + johnsongrass Full season 36%
Mid-May planting One 30 cm-1 of row full season 27% Oliver 1979
Late June planting One 30 cm-1 of row full season 14% Oliver 1979
The Effect of Weed Density 69

greater when warm early-season temperatures Perennial SowthistleSonchus arvensis

favored rapid weed growth.
A field experiment determined that an average of 78
Jerusalem ArtichokeHelianthus tuberosus perennial sowthistle shoots m-2 in 71-cm soybean
rows reduced soybean yield by 49 percent (Zollinger
One paper reports that a density of 1, 2, or 4 tubers
and Kells 1993). In a second, drier year, 96 shoots
m-1 of row reduces soybean yield 31, 59, or 71 per-
m-2 reduced soybean yield 87 percent. One cultiva-
cent (Wyse et al. 1986). Soybeans leaf area and rel-
tion 5 weeks after planting increased crop yield,
ative growth rate were reduced by 2 and 4 tubers m-1
improved seed quality, and decreased perennial
of row and the net assimilation rate was reduced by
sowthistle density.
4 tubers m-1 of row. With 4 tubers m-1 of row for
4, 6, 8, or 20 weeks (full season), soybean yield Pitted MorninggloryIpomoea lacunosa
decreased 9, 10, 38, or 82 percent, respectively. Murdock et al. (1986) used three soybean cultivars
Wyse et al. (1986) concluded that Jerusalem arti- seeded in 30, 61, or 91 cm rows to achieve a uniform
choke should be controlled within 6 weeks after plant population of 323,000 plants ha-1. The narrow-
planting. er row spacings tended to shade the row earlier, and
JimsonweedDatura stramonium some cultivars developed a shading canopy faster. In
one year, the maximum soybean yield was obtained
As the duration of jimsonweed competition with 2 weed-free weeks for 30 and 61 cm rows but
increased, soybean yields decreased, and as jim- no competition was tolerated by soybeans in 91 cm
sonweed emergence was delayed after soybean rows. In a second year, 2 weed-free weeks were tol-
emergence, soybean yield increased (Kirkpatrick et erated by all row spacings and all three cultivars.
al. 1983). As few as 0.3 jimsonweeds m-1 of row Soybeans were grown in conventional 1 m rows and
reduced yield 8 percent and 1.6 weeds m-1 reduced narrow (20 cm) rows with a soybean population of
it 24 percent. Full-season competition from 1.6 23 plants m-2 in 1 m rows and 50 plants
weeds m-1 reduced yield up to 41 percent. Jimson- m-2 in 20 cm rows (Howe and Oliver 1987). Pitted
weed growing in the soybean row had little influ- morningglory density was 3.3, 10, 20, or 40 plants
ence on vegetative characteristics of soybean m-2. Pitted morningglory interfered with soybean
during the first 12 weeks of growth (Henry and growth earlier in conventional rows due to the rapid
Bauman 1991). The influence of a single jimson- increase in its leaf area index and biomass between
weed was not evident until 10 weeks after planting 4 and 8 weeks after planting. The weed was com-
and it then extended 20 cm in the row. By harvest, petitive until soybeans reproductive stages 7 weeks
the influence of a single weed extended 50 cm in after planting, and it decreased soybean yield 17
the row and soybean yield within the area of influ- percent more in a dry year. The yield of narrow-row
ence (1.2 m of row) was reduced 12 percent. Jim- soybeans was equal to or greater than conventional
sonweed that was 60 cm apart in the row reduced rows at all pitted morningglory densities (Howe and
yield 18 percent. Henry and Bauman (1991) report- Oliver 1987). Conventional-row soybean yields
ed that while jimsonweed affected soybean growth, decreased an average of 42 and 81 percent at pitted
the reverse was even more evident. Soybean inter- morningglory densities of 3.3 and 40 plants m-2,
ference with jimsonweed increased steadily during whereas the yield of narrow-row soybeans
the season and reduced the size of jimsonweed decreased only 6 and 62 percent at the same densi-
plants in the row 80 to 93 percent compared to ties. The total seed production of pitted morningglo-
free-standing plants. In contrast, Oliver et al. ry increased as its density increased and was always
(1991) demonstrated that jimsonweed was not par- greater in conventional than in narrow rows. Narrow
ticularly competitive in soybeans but was very rows reduced pitted morningglory seed production
competitive in cotton. Its influence was reduced by an average of 90 and 68 percent when pitted morn-
lack of rain and the fact that soybeans are more ingglory densities were 3.3 or 40 plants m-2. Nors-
competitive than cotton. Soybean yield was worthy and Oliver (2002c) conducted field
reduced 16 percent by 64 jimsonweeds per 12 m-1 experiments to evaluate the role of photosynthetic
of row, whereas cottons was reduced 56 percent by efficiency in the interspecific competition between
the same density. Interspecific interference from soybeans and pitted morningglory planted at 0,
soybeans caused a third more reduction in fresh 10, 16, or 62 plants m-2 in drill-seeded glyphosate-
weight and capsules per plant of jimsonweed than resistant soybeans. Soybeans photosynthetic rate
cotton did (Oliver et al. 1991). was not affected by the weeds density or glyphosate
70 Chapter 5

use 2 weeks after the herbicide was applied. Soy- ties of 95 and 160 shoots m-2 did not affect soybean
beans photosynthetic rate was reduced 12 weeks yield (Young et al. 1982). Interference by a natural
after glyphosate application by 21 and 91 percent stand of quackgrass for 6 or 8 weeks or for the full
when competition was with 62 glyphosate treated or season reduced soybean yield 11, 23, or 33 percent.
untreated pitted morningglory plants m-2. Ten weeds In a separate study, irrigation did not increase the
m-2 did not affect soybeans photosynthetic rate. Pit- yield of quackgrass-free soybeans but the yield of
ted morningglory was not killed by glyphosate, but quackgrass-infested soybeans was increased by irri-
its photosynthetic rate was reduced 2 and 12 weeks gation when soil water was limiting (Young et al.
after glyphosate application by 64 and 80 percent, 1983). They concluded that adequate soil moisture
respectively. Soybean seed yield was not reduced by can reduce quackgrass interference but not eliminate
10 or 16 glyphosate-treated weeds m-2, but a 9 per- it because quackgrass was nearly the same height or
cent loss resulted when there were 62 weeds m-2. taller than soybeans at all stages of soybean devel-
This was partially due to the effect of glyphosate on opment, and it offered significant competition for
the weed and partially due to increased shading by light (Young et al. 1983). Sikkema and Dekker
soybean. Glyphosate did not affect pitted morning- (1987) confirmed quackgrass competition for water
glorys leaf area when the density was 10 or 16 and light, and that competition was partially relieved
plants m-2, but its leaf area was reduced slightly by irrigation. Sikkema and Dekker (1987) also
when the density was 62 weeds m-2. Glyphosate use showed that high levels of phosphorus and potassi-
prevented soybean seed yield reduction, whereas um did not overcome quackgrass interference, but
untreated pitted morningglory reduced soybean its effects were highly variable with a 79 percent
yield 47, 62, or 81 percent at densities of 10, 16, or reduction in soybean yield from full-season compe-
62 plants m-2, respectively (table 5.3). Competitive- tition in one year but only 39 percent in a second
ness of untreated soybeans increased with their year.
seeding rate. There was 22 percent less yield loss
Redroot PigweedAmaranthus retroflexus
when soybeans were planted at 521,000 than at
217,000 plants ha-1. As shown in studies reported above, narrow rows
favor development of soybean leaf area that covers
PoorjoeDiodia teres
the interrow space quickly. Lgre and Schreiber
In a unique experiment, Jordan (1989) used path (1989) were among the first to study plant architec-
analysis to compare growth differences and compet- ture as it influenced interference. They showed that
itiveness of a weedy and a nonweedy (nonagricul- by midseason, redroot pigweed contributed 43 per-
tural) population of poorjoe. Jordan (1989) analyzed cent of the total biomass when soybeans were grown
the establishment rate, early and late growth rates, in 76 cm rows but only 24 percent with 25 cm rows.
growth form, and final growth because each could Soybeans produced two to four times more leaf area
be related to competitiveness. The mean above- than redroot pigweed during the first half of the
ground biomass of the weedy population was rough- growing season, but the advantage was diminished
ly twice that of the nonweedy population whether it with wide rows. Redroot pigweed was 29 percent of
was grown alone or with soybeans. The weedy pop- the total leaf area when soybeans were planted in
ulation of poorjoe had a greater establishment rate 76-cm rows but only 15 percent in 25 cm rows. The
and greater early aboveground growth rate com- leaf area distribution suggested vigorous competi-
pared to the nonweedy population. The final growth tion for light (Lgre and Schreiber 1989).
rate of the two populations was similar whether soy- Redroot pigweed interference in soybeans has not
beans were present or absent. Jordan (1989) postu- been studied as much as has the interference of sev-
lated that there may have been genetic changes in eral other weeds (e.g., common cocklebur, sickle-
the two populations that led to earlier establishment pod, or velvetleaf) because it is not as prevalent in
and faster early growth (characteristic of many soybean growing areas. However, when it interferes
weedy species) but not to an increased tolerance of in soybeans it can be very damaging. In a study of
soybeans or to intraspecific competition. five annual broadleaf species, Shurtleff and Coble
(1985a, b) showed that redroot pigweed at a density
QuackgrassElytrygia repens
of 16 weeds 10 m-1 of row reduced soybean yield
Quackgrass densities of 520 and 910 shoots m-2 loss 22 percent, while the yield reduction due to
reduced soybean yield 19 and 55 percent, but densi- common lambsquarters was 15 percent, common
The Effect of Weed Density 71

ragweed was 12 percent, and sicklepod was 5 per- were a more effective competitor than sorghum as
cent. Eight common cocklebur 10 m-1 of row were demonstrated by the fact that shoot dry weights of
vigorous competitors and reduced yield 11 percent. spleen amaranth were significantly lower when it
Soybean leaf area reductions corresponded with grew with soybeans under the same conditions. Lin-
yield reductions for each weed. The increases in dry ear regression showed that dry weight of the crop
matter and height were slower for all five weeds in was more closely correlated with weed biomass than
this study than for soybeans. The root:shoot ratio of with the number of weeds, and soybeans were more
soybeans was the highest of all plants but, in spite of affected by an increase in weed biomass than
its vigor as a competitor, that of redroot pigweed sorghum (Wulff 1987).
was lowest.
Volunteer CornZea mays
In one of the first multiple species studies to con-
sider time of emergence of the weed relative to the A common (perhaps the most common) rotation in
crop, Cowan et al. (1998) showed that the time of many midwestern fields is corn/soybeans/corn/
emergence of redroot pigweed and barnyardgrass soybeans. Corn frequently becomes a weedy pres-
influenced the amount of soybean yield loss. The ence in soybean fields. Soybean yield is inversely
two weeds were sown together in soybeans when related to volunteer corn clump density, and soybean
the latter was at the cotyledon stage of growth. The yield losses can reach 25 percent with 5,380 clumps
maximum soybean yield loss was 32 to 99 percent of corn per acre. Such clumps may have as many as
depending on the time of emergence. Redroot pig- 10 corn plants (Beckett and Stoller 1988). Corn
weed was more competitive (competitive index of 1 caused soybean yield losses of 2, 6, 12, 19, and 27
on a scale of 0 to 1) than barnyardgrass whose com- percent when it was not controlled for 2, 4, 6, 8,
petitive index was 0.075 to 0.4. or 10 weeks after planting. A 10-corn-plant clump
reduced soybean yield over a radius of 86 cm.
ShattercaneSorghum bicolor
Wild OatAvena fatua
Shattercane has been an important weed in mid-
western U.S. agriculture for many years. It did not Wild oat is common in small grain crops but not
reduce soybean yield if it was removed by 2 weeks usually a major problem in soybean. However, if it
after emergence in one year, and soybeans resisted is present, losses can be significant. Season long
interference for 6 weeks in a second year (Fellows competition of 1, 3, 9, or 30 wild oat m-1 of row
and Roeth 1992). Interference began when shatter- reduced soybean yield an average of 6, 17, 32, or 51
canes height exceeded that of soybeans. Soybean percent (Rathmann and Miller 1981). Thirty wild
yield declined up to 25 percent before the height dif- oats m-1 of row did not affect yield if they were pre-
ferential exceeded 30 cm. This was important sent for 4 weeks after emergence or less. If wild oat
because that height differential was required before was present for 5, 6, 7, 8 weeks or for the full sea-
glyphosate could be applied with a wiper. There was son, yield was reduced 29, 50, 63, 58, or 63 percent.
a direct relationship between shattercane density The effect was more apparent on soybean pods per
and soybean yield that was accurately modeled by a plant and seeds per plant than on seed weight.
rectangular hyperbola. Soybeans height, biomass,
Wild PoinsettiaEuphorbia heterophylla
nodes per stem, pods per stem, pods per node, and
beans per pod all decreased as shattercane density Soybean canopy width was reduced about 10 per-
and the duration of interference increased (Fellows cent beginning after 6 weeks of interference for
and Roeth 1992). distances of 0 to 10 and 10 to 20 cm from the weed.
Soybean dry weights decreased 14 to 38 percent
Spleen AmaranthAmaranthus dubius
within 20 cm of the weed for 12 through 18 weeks
Soybeans were grown in pots with 1, 2, 4, 8, or 16 of interference (Willard et al. 1994). Wild poinset-
spleen amaranth plants in controlled environment tia interference resulted in a 9.5 percent yield
chambers at 31/24 and 25/18C day/night tempera- reduction for the 0 to 10 cm distance from the soy-
tures. Seventeen and 35 days after emergence, bean row compared to 80 to 100 cm. Soybeans
growth of both species had increased with tempera- were an effective competitor with wild poinsettia.
ture. After 35 days, even 1 spleen amaranth per pot Differences in wild poinsettia dry weight when
decreased soybean growth, and the two highest den- growing alone and when growing with soybeans
sities had an equal effect (Wulff 1987). Soybeans occurred after 6 to 8 weeks of interference. In a
72 Chapter 5

year when rainfall was twice that of the previous Bosza, R. C., and L. R. Oliver. 1990. Competitive
year, dry weight of wild poinsettia growing in the mechanisms of common cocklebur (Xanthium stru-
soybean row was reduced 82 percent compared to marium) and soybean (Glycine max) during
the weed growing alone. seedling growth. Weed Sci. 38:344350.
. 1993. Shoot and root interference of common
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The Effect of Weed Density 73

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L. Currey. 1989. The influence of water stress on Klingaman, T. E., and L. R. Oliver. 1994a. Influence
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(Glycine max) and Florida beggarweed (Desmodi- (Glycine max) planting date on weed interference.
um tortuosum). Weed Sci. 37:544551. Weed Sci. 42:6165.
Grymes, C. F., J. L. Griffin, D. J. Boethel, B. R. . 1994b. Palmer amaranth (Amaranthus
Leonard, D. L. Jordan, and J. S. Russin. 1999. Soy- palmeri) interference in soybeans (Glycine max).
bean response to weed interference and defoliation. Weed Sci. 42:523527.
Weed Sci. 47:9094. Knake, E. L. 1972. Effect of shade on giant foxtail.
Harris, T. C., and R. L. Ritter. 1987. Giant green fox- Weed Sci. 20:588592.
tail (Setaria viridis var. major) and fall panicum Krausz, R. F., B. G. Young, G. Kapusta, and J. L.
(Panicum dichotomiflorum) competition in soy- Matthews. 2001. Influence of weed competition
beans (Glycine max). Weed Sci. 35:663668. and herbicides on glyphosate resistant soybean
Harrison, S. K. 1990. Interferences and seed produc- (Glycine max). Weed Technol. 15:530534.
tion by common lambsquarters (Chenopodium Lgre, A., and M. M. Schreiber 1989. Competition
album) in soybeans (Glycine max). Weed Sci. and canopy architecture as affected by soybean
38:113118. (Glycine max), row width and density of redroot
Henry, W. T., and T. T. Bauman. 1991. Interference pigweed (Amaranthus retroflexus). Weed Sci.
between soybean (Glycine max) and jimsonweed 37:8492.
(Datura stramonium) in Indiana. Weed Technol. Lejeune, K. R., J. L. Griffin, D. B. Reynolds, and A.
5:759764. M. Saxton. 1994. Itchgrass (Rottboellia cochinchi-
Higgins, R. A., D. W. Staniforth, and L. R. Pedigo. nensis) interference in soybean (Glycine max).
1984. Effects of weed density and defoliated or Weed Technol. 8:733737.
undefoliated soybeans (Glycine max) on velvetleaf Lindquist, J. L., B. D. Maxwell, D. D. Buhler, and J.
(Abutilon theophrasti) development. Weed Sci. L. Gunsolus. 1995. Velvetleaf (Abutilon
32:511519. theophrasti) recruitment, survival, seed production,
Howe, O. W., III, and L. R. Oliver. 1987. Influence of and interference in soybean (Glycine max). Weed
soybean (Glycine max) row spacing on pitted morn- Sci. 43:226232.
ingglory (Ipomoea lacunosa) interference. Weed McWhorter, C. G., and G. L. Sciumbato. 1988.
Sci. 35:185193. Effects of row spacing, benomyl, and duration of
Irons, S. M., and O. C. Burnside. 1982. Competitive sicklepod (Cassia obtusifolia) interference on soy-
and allelopathic effects of sunflower (Helianthus bean (Glycine max) yields. Weed Sci. 36:254259.
annuus). Weed Sci. 30:372377. McWhorter, C. G., and J. M. Anderson. 1993. Effects
Jackson, L. A., G. Kapusta, and D. J. Schutte-Mason. of johnsongrass (Sorghum halepense), hemp sesba-
1985. Effect of duration and type of natural weed nia (Sesbania exaltata), and delayed harvest on
infestation on soybean yield. Agron. J. 77:725729. soybeans. Weed Technol. 7:355360.
74 Chapter 5

Monks, D. W., and L. R. Oliver. 1988. Interactions Parker, M. B., H. W. Marchant, and B. J. Mullinix, Jr.
between soybean (Glycine max) cultivars and 1981. Date of planting and row spacing effects on
selected weeds. Weed Sci. 36:770774. four soybean cultivars. Agron. J. 73:759762.
Mortensen, D. A., and H. D. Coble. 1989. The influ- Patterson, D. T. 1986. Interference of five broadleaf
ence of soil water content on common cocklebur weeds in soybeans. Weed Sci. Soc. America. Abst.
(Xanthium strumarium) interference in soybeans No. 163, p. 59.
(Glycine max). Weed Sci. 37:7683. Patterson, D. T., and E. P. Flint. 1983. Comparative
Mosier, D. G., and L. R. Oliver. 1995a. Common water relations, photosynthesis, and growth of soy-
cocklebur (Xanthium strumarium) and entireleaf bean (Glycine max) and seven associated weeds.
morningglory (Ipomoea hederacea var. Weed Sci. 31:318323.
integriscula) interference in soybeans (Glycine Patterson, M. G., R. H. Walker, D. L. Colvin, G.
max). Weed Sci. 43:239246. Wehtje, and J. A. McGuire. 1988. Comparison of
. 1995b. Soybean (Glycine max) interference soybean (Glycine max)weed interference from
on common cocklebur (Xanthium strumarium) and large and small plots. Weed Sci. 36:836839.
entireleaf morningglory (Ipomoea hederacea var. Quakenbush, L. S., and R. N. Andersen. 1984. Effect
integriuscula). Weed Sci. 43:402409. of soybean (Glycine max) interference on Eastern
Mulugeta, D., and C. M. Boerboom. 2000. Critical black nightshade (Solanum ptycanthum). Weed Sci.
time of weed removal in glyphosate-resistant 32:638645.
Glycine max. Weed Sci. 48:3542. Rathmann, D. P., and S. D. Miller. 1981. Wild oat
Munger, P. H., J. M. Chandler, J. T. Cothern. 1987. (Avena fatua) competition in soybean (Glycine
Effect of water stress on photosynthetic parameters max). Weed Sci. 29:410414.
of soybean (Glycine max) and velvetleaf (Abutilon Regnier, E. E., and S. K. Harrison. 1993. Community
theophrasti). Weed Sci. 35:1521. responses of common cocklebur (Xanthium stru-
Munger, P. H., J. M. Chandler, J. T. Cothern, and F. marium) and velvetleaf (Abutilon theophrasti) to
M. Honsi. 1987. Soybean (Glycine max) partial shading. Weed Sci. 41:541547.
velvetleaf (Abutilon theophrasti) interspecific Regnier, E. E., and E. W. Stoller. 1989. The effects of
competition. Weed Sci. 35:647653. soybean (Glycine max) interference on the canopy
Murdock. E. C., P. A. Banks, and J. E. Toler. 1986. architecture of common cocklebur (Xanthium stru-
Shade development effects on pitted morningglory marium), jimsonweed (Datura stramonium), and
(Ipomoea lacunosa) interference in soybeans. Weed velvetleaf (Abutilon theophrasti). Weed Sci.
Sci. 34:711717. 37:187195.
Murphy, T. R., and B. J. Gossett. 1981. Influence of Regnier, E. E., E. W. Stoller, and E. D. Nafziger.
shading by soybeans (Glycine max) on weed sup- 1989. Common cocklebur (Xanthium strumarium)
pression. Weed Sci. 29:610615. root and shoot interference in soybeans (Glycine
Norsworthy, J. K., and L. R. Oliver. 2002a. Effect max). Weed Sci. 37:308313.
of irrigation, soybean (Glycine max) density, Rose, S. J., O. C. Burnside, J. E. Specht, and B. A.
and glyphosate on hemp sesbania (Sesbania Swisher. 1984. Competition and allelopathy between
exaltata) and pitted morningglory (Ipomoea soybeans and weeds. Agron. J. 76:523528.
lacunosa) interference in soybean. Weed Tech- Rushing, G. S., and L. R. Oliver. 1998. Influence of
nol. 16:717. planting date on common cocklebur (Xanthium
. 2002b. Hemp sesbania interference in drill- strumarium) interference in early-maturing soybean
seeded glyphosate-resistant soybean. Weed Sci. (Glycine max). Weed Sci. 46:99104.
50:3441. Scott, H. D., and R. D. Geddes. 1979. Plant water
. 2002c. Pitted morningglory interference in stress of soybean (Glycine max) and common cock-
drill-seeded glyphosate-resistant soybean. Weed Sci. lebur (Xanthium pensylvanicum): A comparison
50:2633. under field conditions. Weed Sci. 27:285289.
Oliver. L. R. 1979. Influence of soybean (Glycine Schultz, M. E., and O. C. Burnside. 1979. Distribu-
max) planting date on velvetleaf (Abutilon tion, competition and phenology of hemp dogbane
theophrasti) competition. Weed Sci. 27:183188. (Apocynum cannibinum) in Nebraska. Weed Sci.
Oliver, L. R., J. M. Chandler, and G. A. Buchanan. 27:565570.
1991. Influence of geographic region on jimson- Shaw, D. R., A. Rankins, Jr., and J. T. Ruscoe. 1997.
weed (Datura stramonium) interference in soybeans Sicklepod (Senna obtusifolia) interference with
(Glycine max) and cotton (Gossypium hirsutum). soybean (Glycine max) cultivars following herbi-
Weed Sci. 39:585589. cide treatments. Weed Technol. 11:510514.
The Effect of Weed Density 75

Shaw, D. R., S. A. Bruff, and C. A. Smith. 1991. Walker, R. H., M. G. Patterson, E. Hauser, D. J. Isen-
Effect of soybean (Glycine max) row spacing on hour, J. W. Todd, and G. A. Buchanan. 1984.
chemical control of sicklepod (Cassia obtusifolia). Effects of insecticide, weed-free period, and row
Weed Technol. 5:286290. spacing on soybean (Glycine max) and sicklepod
Shilling, D. G., B. J. Brecke, C. Hiebsch, and G. Mac- (Cassia obtusifolia) growth. Weed Sci. 32:702706.
Donald. 1995. Effect of soybean (Glycine max) cul- Webster, T. M., J. Cardina, and S. J. Woods. 2000.
tivar, tillage, and rye (Secale cereale) mulch on Apocynum cannabinum interference in no-till
sicklepod (Senna obtusifolia). Weed Technol. Glycine max. Weed Sci. 48:716719.
9:339342. Webster, T. M., M. Loux, E. E. Regnier, and S. K.
Shurtleff, J. L., and H. D. Coble, 1985a. Interference Harrison. 1994. Giant ragweed (Ambrosia trifida)
of certain broadleaf weed species in soybeans canopy architecture and interference studies in soy-
(Glycine max). Weed Sci. 33:654657. bean (Glycine max). Weed Technol. 8:559564.
. 1985b. The interaction of soybean (Glycine Willard, T. S., J. L. Griffin, D. B. Reynolds, and A. M.
max) and five weed species in the greenhouse. Saxton. 1994. Interference of wild poinsettia
Weed Sci. 33:669672. (Euphorbia heterophylla) with soybean (Glycine
Sikkema, P. H., and J. Dekker. 1987. Use of infared max). Weed Technol. 8:679683.
thermometry in determining critical stress periods Williams, C. S., and R. M. Hayes. 1984. Johnsongrass
induced by quackgrass (Agropyron repens) in soy- (Sorghum halepense) competition in soybeans
bean (Glycine max). Weed Sci. 35:784791. (Glycine max). Weed Sci. 32:498501.
Sims, B. D., and L. R. Oliver. 1990. Mutual influ- Wulff, R. D. 1987. Growth responses of soybean
ences of seedling johnsongrass (Sorghum (Glycine max) and sorghum (Sorghum bicolor) to
halepense), sicklepod (Cassia obtusifolia), and soy- an increase in density of Amaranthus dubius L. at
bean (Glycine max). Weed Sci. 38:139147. two temperatures. Weed Res. 27:7986.
Smith, J. E., and P. W. Jordan. 1993. Sicklepod (Cas- Wyse, D. L., F. L. Young, and R. J. Jones. 1986. Influ-
sia obtusifolia) shoot structure as affected by soy- ence of Jerusalem artichoke (Helianthus tuberosus)
bean (Glycine max) interference. Weed Sci. density and duration of interference on soybean
41:7581. (Glycine max) growth and yield. Weed Sci.
Smith, R. J., Jr. 1988. Weed thresholds in southern 34:243247.
U.S. rice. Weed Technol. 2:232241. Yelverton, F. C., and H. D. Coble. 1991. Narrow row
Stoller, E. W., S. K. Harrison, L. M. Wax, E. E. Reg- spacing and canopy formation reduces weed resur-
nier, and E. D. Nafziger. 1987. Weed interference in gence in soybean (Glycine max). Weed Technol.
soybeans (Glycine max) Rev. Weed Sci. 3:155181. 5:169174.
Stoller, E. W., and R. A. Myers. 1989a. Effects of Young, F. L., D. L. Wyse, and R. J. Jones. 1982. Influ-
shading and soybean (Glycine max L.) Interference ence of quackgrass (Agropyron repens) density and
on Solanum ptycanthum Dun. (eastern black night- duration of interference in soybeans (Glycine max).
shade) growth and development. Weed Res. Weed Sci. 30:614619.
29:307316. . 1983. Effect of irrigation on quackgrass
. 1989b. Response of soybeans (Glycine max) (Agropyron repens) interference in soybeans
and four broadleaf weeds to reduced irradiance. (Glycine max). Weed Sci. 31:720727.
Weed Sci. 37:570574. Zimdahl, R. L. 1980. Weed-crop competition: A
Toler, J. E., J. B. Guice, and E. C. Murdock. 1996. review. Corvallis, OR: Int. Plant Prot. Center, Ore-
Interference between johnsongrass (Sorghum gon State University.
halepense), smooth pigweed (Amaranthus Zollinger, R. K., and J. J. Kells. 1993. Perennial
hybridus), and soybean (Glycine max). Weed Sci. sowthistle (Sonchus arvensis) interference in soy-
44:331338. bean (Glycine max) and dry edible bean (Phaseolus
United Nations Food and Agriculture Organization. vulgaris). Weed Technol. 7:5257.
2000. Production yearbook 54: 7677.
Van Acker, R. C., S. F. Wiese, and C. J. Swanton. SUGARBEETBETA VULGARIS L.
1993. Influence of interference from a mixed weed
species stand on soybean [Glycine max (L.) Merr.] Sugarcane and sugarbeets are the worlds primary
growth. Can J. Plant Sci. 73:12931304. sources of sugar, a completely nonessential but quite
Vitta, J. I., and E. H. Satorre. 1999. Validation of a enjoyable part of the human diet. Sugar is a crop that
weed:crop competition model. Weed Res. has changed the course of human history in undesir-
39:259269. able ways (Hobhouse 1986).
76 Chapter 5

The response of a semiprostrate and an erect sug- measure (i.e., degree days) instead of days. The
arbeet cultivar to nitrogen and to wild mustard and wisdom of this suggestion has been recognized by
common lambsquarters was not identical. Timing of many. Kropff et al. (1992) also reported that mor-
nitrogen fertilization did not influence crop biomass phological characteristics such as relative growth
or yield and crop quality of the weed-free crop. rate of leaf area and height are the primary determi-
Early nitrogen application resulted in opposite nants of competitive ability whereas physiological
effects with the two weeds: higher crop biomass traits such as maximum rate of photosynthesis are
reduction in the presence of wild mustard and lower less significant. In competition, morphology (size)
crop biomass in the presence of common lambs- is more important than physiology in most cases.
quarters. Root and sucrose yield responded similar- Sugarbeet root yield was reduced by competition
ly to weed competition. There was no difference in from all densities of wild oat and wild mustard alone
cultivar response to weed competition. The crop was or in combination in Wyoming (Mesbah et al. 1995).
favored by late nitrogen application if wild mustard Root yield reduction was less than additive when the
was present but by early nitrogen application when two weeds were mixed. Root yield decreased as the
common lambsquarters was the competing species duration of interference increased but percent
(Paolini et al. 1999). sucrose was not affected. The maximum time a mix-
When wild mustard or perennial ryegrass were ture of 0.8 wild mustard and 1 wild oat m-1 of row
grown in the sugarbeet row or 2, 4, or 8 cm from the can be present without yield loss is 1.6 weeks after
row, sugarbeet yield increased when the weeds were sugarbeet emergence.
further from the crop row (Heisel et al. 2002). When Similar results were found for mixed densities
the distance increased from 2 to 8 cm, sugarbeet and durations of competition from kochia and green
yield increased 20 percent, regardless of the weed foxtail (Mesbah et al. 1994). With these species,
species. The number of neighbors described a sig- reductions in yield were additive at low densities but
moidal yield decline of single sugarbeet plants. were less than additive (see above for wild oat and
Results of image analysis (analysis of leaf cover) wild mustard) at high densities. Because sugarbeets
showed that about 33 g of sugarbeet yield was lost were irrigated and fertilizer was applied to optimize
in October/November for each percent of projected yield, the authors concluded that both weeds com-
leaf cover of weeds in May (Heisel et al. 2002). peted primarily for light. The lowest densities of
Kropff and Spitters (1992) proposed an ecophysi- kochia and green foxtail were 0.3 and 0.06 plants
ological simulation model based on how the sugar- m-1 of row, respectively. The minimum time that 0.5
beet and common lambsquarters use light and water kochia and 3 green foxtail m-1 of row can interfere
for dry matter production. The distribution of the with sugarbeets without economic loss of root yield
leaf area of the competing species over the sugarbeet is about 3.5 weeks after sugarbeet emergence (Mes-
canopy was used to determine absorbed radiation in bah et al. 1994), a longer time than was true for wild
relation to canopy height. The CO2 assimilation light oats and wild mustard (Mesbah et al. 1994).
response of individual leaves was used to calculate Interference of barnyardgrass with 10 or more
the canopys CO2 assimilation profile. A daily CO2 plants m-1 of row caused more than 80 percent root
assimilation rate was calculated for each species. yield decrease in spring-planted sugarbeets in Cali-
Soil moisture and drought were considered. Subse- fornia (Norris 1992). Yield loss was only 5 to 20 per-
quent work showed that 98 percent of the variation cent when there was 1 weed in every 2 to 3 m of row.
in yield loss (range -6 to 96 percent) was explained The economic threshold density was about one
by the simulation model (Kropff et al. 1992). The barnyardgrass plant 5 to 20 m-1 of row. If barnyard-
primary factor responsible for difference in yield grass was present at densities less than 1 m-1 of row,
loss between experiments was the number of days sugarbeets experienced only interspecific competi-
between crop and weed emergence (0 to 31 days). tion and the weeds had no measurable effect. How-
Water shortage only influenced the competitive abil- ever, because barnyardgrass is a prolific seed
ity of the weeds when they were shorter than the producer, one weed 10 m-1 of row produced between
crop. Temperature in the period between crop and 4,000 and 20,000 seeds m-1 of row (Norris 1992).
weed emergence was also an important determinant The weeds presence in future crops is assured even
of competitive ability. Kropff et al. (1992) recom- though the damage to the present crop is minimal.
mended that the time between crop and weed emer- A series of experiments was conducted by
gence should be expressed as a developmental Schweizer and colleagues in Colorado to determine
The Effect of Weed Density 77

the interference of different weed species in sugar- Kropff, M. J., and C. J. T. Spitters. 1992. An ecophys-
beets. When equal densities of kochia, redroot pig- iological model for interspecific competition
weed, and common lambsquarters were present at applied to the influence of Chenopodium album L.
total densities of 3, 6, 12, 18, or 24 weeds in 30 m of on sugar beet. I. Model description and parameteri-
row, sugarbeet root yields decreased not at all, 13, zation. Weed Res. 32:437450.
24, 33, or 39 percent, respectively (Schweizer Kropff, M. J., and C. J. T. Spitters, B. J. Schneiders,
1981). A linear equation predicted root yield loss W. Joenje, and W. de Groots. 1992. An ecophysio-
with increasing weed density. The actual yield loss logical model for interspecific competition applied
was always less than the predicted loss because the to the influence of Chenopodium album L. on sugar
beet. II. Model evaluation. Weed Res. 32:451464.
growth of weeds to which herbicides were applied
Mesbah, A., S. D. Miller, K. J. Fornstrom, and D. E.
was suppressed, but they remained competitive.
Legg. 1994. Kochia (Kochia scoparia) and green
Powell amaranth at 6, 12, 18, or 24 in 30 m of row
foxtail (Setaria viridis) interference in sugarbeets
was less competitive than the weeds above (Beta vulgaris). Weed Technol. 8:754759.
(Schweizer and Lauridson 1985) and reduced yield . 1995. Wild mustard (Brassica kaber) and
7, 13, 23, or 24 percent. The minimum number of wild oat (Avena fatua) interference in sugarbeets
Powell amaranth required in 30 m of row to reduce (Beta vulgaris). Weed Technol. 9:4952.
yield was 9 in one year and 11 in a second year of Norris, R. F. 1992. Case history for weed competition/
study. Common lambsquarters at 6, 12, 18, or 24 in population ecology in sugarbeets (Beta vulgaris).
30 m of row reduced yield 1113, 2729, 3738, or Weed Technol. 6:220227.
4648 percent, respectively, in 2 years of study. It Paolini, R., M. Principi, R. J. Froud-Williams, S. Del
was more competitive than Powell amaranth Puglia, and E. Binacardi. 1999. Competition
because the minimum number of weeds required to between sugarbeet and Sinapis arvensis and
reduce yield was 4 in one year and 6 in a second Chenopodium album as affected by timing of nitro-
year of study (Schweizer 1983). Common sunflower gen fertilization. Weed Res. 39:425440.
was more competitive than velvetleaf when densi- Schweizer, E. E. 1981. Broadleaf weed interference in
ties identical to those reported above were used. sugarbeets (Beta vulgaris). Weed Sci. 29:128133.
Yield reduction from common sunflower was 40, . 1983. Common lambsquarters (Chenopodium
52, 67, or 73 percent, whereas those from velvetleaf album) interference in sugarbeets. Weed Sci.
were only 14, 17, 25, or 30 percent for the same den- 31:58.
sity (Schweizer and Bridge 1992). When both weeds Schweizer, E. E., and L. D. Bridge. 1982. Sunflower
competed with sugarbeet, yield losses were interme- (Helianthus annuus) and velvetleaf (Abutilon
theophrasti) interference in sugarbeets (Beta vul-
diate between those for either weed alone with sug-
garis). Weed Sci. 30:514519.
arbeet, indicating interspecific competition between
Schweizer, E. E., and T. C. Lauridson. 1985. Powell
the weeds. The minimum number of weeds required
amaranth (Amaranthus powellii) interference in
to reduce sugarbeet yield was 1 common sunflower, sugarbeets (Beta vulgaris). Weed Sci. 33:518520.
9 to 12 velvetleaf, and 2 to 7 when the weeds were
mixed. Common sunflower was a more effective
competitor because of its rapid early growth and
large size late in the season. When sugarbeets were
harvested, common sunflower averaged 240 cm tall, Sugarcane is an important cash crop in many parts
kochia was 157 cm, velvetleaf was 150, and sugar- of the world. In spite of its importance but because
beets only 50 cm (Schweizer and Bridge 1982). of its long growing period, dense foliage, height,
and ready availability of several selective herbicides,
Literature Cited weed competition has not been studied as much as it
Heisel, T., C. Andreasen, and S. Christensen. 2002. has in most annual crops.
Sugarbeet yield response to competition from Itchgrass germinates throughout the cane growing
Sinapis arvensis or Lolium perenne growing at season and is a vigorous competitor. There was no
three different distances from the beet and removed difference in the sugarcane population, the yield of
at various times during early growth. Weed Res. cane, or the sugar yield when itchgrass was allowed
42:406413. to interfere for the entire season, from early-season
Hobhouse, H. 1986. Seeds of change: Five plants that emergence until the last cultivation in mid-June, or
transformed mankind. New York: Harper & Row. from late-season (the last cultivation) until sugarcane
78 Chapter 5

harvest. Full-season interference reduced sugarcane VEGETABLES

population and cane and sugar yield an average of 34,
BeanPhaseolus spp. and Vicia spp.
42, and 43 percent, respectively, compared with no
interference in Louisiana (Lencse and Griffin 1991). The sensitivity of beans to weed competition is illus-
Autumn-planted sugarcane seeded with 1 itchgrass in trated by work with redstem filaree (Blackshaw and
30.5 m of row (= 1.8 itchgrass m-2)in early March lost Harker 1998). Maximum yield reduction from 100
an average of 7 percent of sugar yield after 30 days of to 200 weeds m-2 for wheat was 36 percent and
interference (Millhollen 1992). After 30 days, itch- oilseed rape was 37 percent. For beans, the maxi-
grass biomass rangedfrom 200 to 2,700 kg ha-1 but mum reduction was 82 percent and only pea was
after 60 days it had increased to 1,400 to 2,900 kg more sensitive to competition with a loss of 92 per-
ha-1 and sugar yield was reduced 17 percent. When cent. Three weeks of competition from redstem fila-
itchgrass interfered until harvest (180 days), sugar ree was sufficient to reduce bean yield and the mean
yield was reduced 19 percent, and one must conclude yield reduction for bean per week of competition
that early interference is most damaging. The weeds was 6.3 percent, higher than the other three crops.
primary effect was a reduction in sugarcane stalk den- Burnside et al. (1998) reported that the critical
sity. When itchgrass was removed in the crops sec- period for weed control (see chapter 6) in dry beans
ond year (the ratoon crop) on May 1, June 1, or in Nebraska was 3 to 5 or 6 weeks after planting.
November 15, yield was reduced 3, 11, or 72 percent Weed-removal timing had little effect on dry bean
compared to a weed-free crop. However, sugarcane stand or 100 seed weight of harvested seed. Also in
stand and yield recovered almost completely when Nebraska, Wilson (1993) reported that wild proso
the crop was maintained weed free in the second year millet was able to reduce bean yield between 12 and
following full-season itchgrass interference in the 31 percent at a density of 10 weeds m-2. The rectan-
first crop year (Millhollen 1992). gular hyperbolic model predicted the weeds effect
Losses up to 40 percent of cane yield were caused on bean yield as weed density increased. Consistent
by natural weed populations in the Sudan (Ibrahim with the work of Burnside et al. (1998), Wilson
1984). Weed competition decreased millable stalks found a weed-free period of 4 weeks was sufficient
(32 percent), stalk thickness (15 percent), and the to prevent yield loss.
number of nodes per stalk (14 percent). Four hand As few as two hairy nightshade plants in a meter
weedings were no better than three weedings 3, 6, of row reduced bean (red bean) yield an average of
and 9 weeks after cane planting. 13 percent (Blackshaw 1991). If the weed was pre-
Johnsongrass, a tall perennial grass, decreased sent during the first 3 weeks after bean emergence,
cane yield 36 percent and sugar yield 31 percent bean yield was depressed. Dependent on the length
compared to weed-free plots in Louisiana (Ali et al. of the growing season, 6 to 9 weeks of weed-free
1986). The weeds greatest effect was on cane den- maintenance were required after emergence to pre-
sity (i.e., stalks per plot) and effects were only vent hairy nightshade from producing viable seed
observed when johnsongrass exceeded 15 to 35 per- before frost killed the weed. Up to 9 weeks of weed-
cent of total plant density. free maintenance were required to prevent bean
yield loss.
Literature Cited As reported in chapter 6, when the critical period
for white bean (also Phaseolus spp.) was defined as
Ali, A. D., T. E. Reagan, L. M. Kitchen, and J. L.
the beginning of the crop stage of growth when
Flynn. 1986. Effects of johnsongrass (Sorghum
halepense) density on sugarcane (Saccharum offici-
weed presence reduced yield by 3 percent and end-
narum) yield. Weed Sci. 34:381383. ing at the crop growth stage to which the crop had to
Ibrahim, A. A. S. 1984. Weed competition and control be weed free to prevent a 3 percent yield loss, the
in sugarcane. Weed Res. 24:227232. critical period extended from the second trifoliate to
Lencse, R. J., and J. L. Griffin. 1991. Itchgrass (Rot- the first flower stage of growth for all cultivars. The
tboellia cochinchinensis) interference in sugarcane beginning of the critical period corresponded with
(Saccharum sp.). Weed Technol. 5:396399. the beginning of a rapid increase in total weed bio-
Millhollen, R. W. 1992. Effect of itchgrass (Rottboel- mass (Woolley et al. 1993).
lia cochinchinensis) interference on growth and Bean cultivars varied in their competitive ability
yield of sugarcane (Saccharum officinarum). Weed against a natural population of annual weeds in
Sci. 40:4853. Ontario, Canada. Uncontrolled weed populations
The Effect of Weed Density 79

reduced bean yield as much as 70 percent. Two cul- cent compared to conventional 91 cm rows (Teas-
tivars reduced weed growth 10 to 35 percent com- dale and Frank 1983). If the weeds were controlled
pared to a third cultivar (Malik et al. 1993). All for the first half of the growing season, 15 to 35 cm
cultivars were more competitive in medium-width rows suppressed weed growth 82 percent compared
(46 cm) and narrow (23 cm) rows than in wide (the to 91 cm rows. The effect of 46 cm rows was vari-
traditional width) 69 cm rows. Neither the cultivar, able. Narrow rows suppressed weeds because the
row spacing, nor planting density alone had a sig- bean canopy closed sooner. Beans in 15, 25, 36, or
nificant effect on weed density. However, the com- 46 cm rows had similar yields that were an average
bination of cultivar, row spacing, and planting of 23 percent higher than beans in 91 cm rows.
density that maximized beans leaf area index also In fields where horsenettle had been grown for 3
minimized weed biomass (Malik et al. 1993). Each years before beans were planted, bean yield was
kg ha-1 increase in weed biomass increased white reduced 48 percent in one and 65 percent in another
beans yield loss by 0.38 kg ha-1. year (Frank 1990). Horsenettle that had been estab-
The time of common ragweed emergence and the lished for only 1 year reduced yield 18 to 20 percent.
weeds density affected bean yield at all locations in When horsenettle was 15, 30, or 60 cm from the
work by Chikoye et al. (1995). When 1.5 common bean row, yield was reduced 43, 29, and 15 percent
ragweed plants emerged in a 1 meter of row at in one year and 38, 26, or 11 percent in a second
beans VE (seedling) stage, seed yield loss was 10 to year, illustrating the vigor of horsenettle competi-
22 percent. When emergence of the same density tion.
was delayed until beans V3 (second trifoliate) Eight horsenettle plants in 4.6 m of row reduced
stage, yield loss was only 4 to 9 percent. The time bean yield 36 percent and 16 reduced it 55 percent.
that common ragweed emerged was more important If beans were planted in 15 cm rows, horsenettle
than its density as a determinant of bean yield. fruit production was reduced 16 percent from that
Another of the few studies of interference of para- produced in 60 cm rows.
sitic weeds was done, as many have been, in Spain. Intraspecific competition was always more
Crenate broomrape growth was weakly negatively severe than interspecific competition when red kid-
correlated with the final shoot height or number of ney beans (Phaseolus sp.) competed with black
shoots of broad bean (Mesa-Garcia and Garcia-Torres nightshade or barnyardgrass in California (Fenni-
1984). The primary effect of crenate broomrape was more et al. 1984). By 47 days after planting, barn-
on the number of bean pods. Second, the number of yardgrass and bean both reduced bean biomass and
seeds in each pod was reduced when competition yield. Beans were better competitors than either
occurred at late crop growth stages, when pods had weed.
already developed. The average of 4 crenate broom- In contrast, common cocklebur reduced bean
rape plants parasitizing 1 broad bean plant reduced yield 8 to 44 percent in one year and 2 to 55 percent
seed yield 50 percent. A second study, in northwest in a second year by full-season competition from
Syria (Manschadi et al. 1997), showed that the num- densities between 0.5 and 8 weeds m-2. In one year,
ber of crenate broomrape attachments in one faba 1 weed m-2 was the damage threshold and 4 m-2 was
bean genotype was positively correlated with plant the threshold in the second year. Snap beans could
density. However, faba bean planting density had no compete effectively only until the unifoliate stage.
significant effect on crenate broomrape in either cul- The critical duration of interference for common
tivar. Resistance in a second genotype was due to cocklebur that emerged with snap beans was
three characteristics of the genotype. First it had between emergence and the full-bloom stage of snap
reduced plant vigor and reduced root-length density. beans (Neary and Majek 1990).
Second, just before or just after the parasites pene- One study of interference of weeds in lima
tration into the hosts root, host cell necrosis occurred beans was found. Sicklepod was studied at densi-
that effectively created a barrier to further penetra- ties between 2.7 and 43.1 weeds m-2 for 0, 2, 4, 6,
tion. Finally, the genotype had early flowering and 8, or 10 weeks after planting (Glaze and Mullinix
pod set and thus matured more rapidly, which 1984). Lima bean yield was inversely related to
reduced the effects of the parasite observed on faba sicklepod density. In general, 2.7 and 10.8 sickle-
bean cultivars that were in the field longer. pod m-2 did not reduce lima bean yield. Bean yield
When weeds emerged with beans, crop row spac- decreased as the duration of competition exceeded
ings of 15 to 36 cm suppressed weed growth 18 per- 6 weeks.
80 Chapter 5

Literature Cited LentilLens culinaris L. and chickpea

Cicer arietinum L.
Blackshaw, R. E. 1991. Hairy nightshade (Solanum
sarrachoides) interference in dry beans (Phaseolus Competition from four weeds during the first 30
vulgaris). Weed Sci. 39:4853. days after planting decreased lentil grain yield an
Blackshaw, R. E., and K. N. Harker. 1998. Erodium average of 17 percent. Losses increased with time of
cicutarium density and duration of interference competition up to 69 percent for full-season compe-
effects on yield of wheat, oilseed rape, pea, and dry tition (Singh and Singh 1990). Yield increased with
bean. Weed Res. 38:5562. the length of an early weed-free period up to 60 days
Burnside, O. C., M. J. Wiens, B. H. Holder, S. Weis- after planting, after which the crop effectively sup-
berg, E. A. Ristau, M. M. Johnson, and J. H.
pressed weed growth.
Cameron. 1998. Critical periods for weed control in
In Jordan, competition from a natural weed stand
dry beans (Phaseolus vulgaris). Weed Sci.
with winter-planted, rain-fed lentil and chickpea
was studied over 2 years (Al-Thahabi et al. 1994).
Chikoye, D., S. F. Wiese, and C. J. Swanton. 1995.
Influence of common ragweed (Ambrosia artemisi-
Chickpea seed yield was reduced an average of 81
ifolia) time of emergence and density on white percent, and straw yield declined 63 percent after
bean (Phaseolus vulgaris). Weed Sci. 48:375380. full-season competition. The critical period for com-
Fennimore, S. A., L. W. Mitich, and S. R. Radosevich. petition was 35 to 49 days after chickpea emer-
1984. Interference among bean (Phaseolus gence. Lentil seed yield decreased 63 percent and
vulgaris), Barnyardgrass (Echinochloa crus-galli), straw yield declined 55 percent after full-season
and black nightshade (Solanum nigrum). Weed Sci. competition. The critical period for lentil was 49 to
32:336342. 56 days after emergence. The critical period for both
Frank, J. R. 1990. Influence of horsenettle (Solanum crops occurred when they were in an advanced
carolinense) on snapbean (Phaseolus vulgaris). stage of vegetative growth but before flowering.
Weed Sci. 38:220223. There was a significant loss of biomass and yield
Glaze, N. C., and B. G. Mullinix, Jr. 1984. Competi- in 3 of 4 years when round-leaved mallow compet-
tive effects of sicklepod on lima beans. Weed Sci. ed with lentils. A two-variable model that consid-
32:13. ered early-season crop density loss and
Malik, V. S., C. J. Swanton, and T. E. Michaels. 1993. round-leaved mallow density best accounted for
Interaction of white bean (Phaseolus vulgaris L.) variation in lentil and wheat yield. Losses could be
cultivars, row spacing, and seeding density with up to 100 percent in lentil from full-season compe-
annual weeds. Weed Sci. 41:6268. tition (Makowski 1995). In lentil, 200 round-leaved
Manschadi, A. M., J. Sauerborn, J. Kroschel, and M. mallow m-2 had a biomass of 200 to 1,000 g m-2,
C. Saxena. 1997. Effect of plant density on grain
nearly twice that of the weed in wheat.
yield, root-length density and Orobanche crenata
An infestation of 32 or 65 wild oats m-2 for 5
infestation in two faba bean genotypes. Weed Res.
weeks after lentil emergence did not reduce lentil
Mesa-Garcia, J., and L. Garcia-Torres. 1984. A com- yield (Curran et al. 1987). However, 32 wild oats
petition index for Orobanche crenata Forsk. effects m-2 reduced yield 32 percent if present for 7 weeks
on broad bean (Vicia faba). Weed Res. 24:37938 and 49 percent if present to harvest (11 weeks). Wild
Neary, P. E., and B. A. Majek. 1990. Common cockle- oats decreased grain yield 42 and 61 for 7 or 11
bur (Xanthium strumarium) interference in snap weeks when present at 65 m-2.
beans (Phaseolus vulgaris). Weed Technol. Literature Cited
Teasdale, J. R., and J. R. Frank. 1983. Effect of row Al-Thahabi, S. A., J. Z. Yasin, B. E. Abu-Irmaileh, N.
spacing on weed competition with snap beans I. Haddad, and M. C. Saxena. 1994. Effect of weed
(Phaseolus vulgaris). Weed Sci. 31:8185. removal on productivity of chickpea (Cicer ariet-
Wilson, R. G. 1993. Wild proso millet (Panicum mili- inum) and lentil (Lens culinaris Med.) in a Mediter-
aceum) interference in dry beans (Phaseolus vul- ranean environment. J. Agron. and Crop Sci.
garis). Weed Sci. 41:607610. 172:333341.
Woolley, B. L., T. E. Michaels, M. R. Hall, and C. J. Curran, W. S., L. A. Morrow, and R. E. Whitesides.
Swanton. 1993. The critical period of weed control 1987. Lentil (Lens culinaris) yield as influenced by
in white bean (Phaseolus vulgaris). Weed Sci. duration of wild oat (Avena fatua) interference.
41:180184. Weed Sci. 35:669672.
The Effect of Weed Density 81

Makowski, R. M. D. 1995. Round-leaved mallow sensitive of the four crops studied with a maximum
(Malva pusilla) interference in spring wheat yield reduction of 92 percent. Three weeks of com-
(Triticum aestivum) and lentil (Lens culinaris) in petition from redstem filaree was sufficient to
Saskatchewan. Weed Sci. 43:381388. reduce bean yield; the mean pea yield reduction per
Singh, G., and D. Singh. 1990. Weed competition week of competition was 3.6 percent, lower than for
studies in lentil (Lens culinaris Medic.). Indian J. bean.
Weed Sci. 22:16. Vined pea yield ha-1 was reduced by weeds by a
OnionAllium cepa L. constant amount across a range of densities (Lawson
1983). In general, weeds had effects similar to the
A density of 18 London rocket m-2 reduced onion effects of increasing pea crop density, but without
yield 6 weeks after planting (Menges and Tamez the added contribution the extra pea plants made to
1981b). Onion, a noncompetitive crop, also lost yield. Higher density pea crops suppressed weeds
yield when 360 common sunflowers m-2 interfered effectively but were as vulnerable to yield loss as
for 6 weeks after emergence and when the same crops with lower density except the adverse effects
weed was present at a density of 50 or 5 m-2 for 12 of weeds were diminished by the interspecific com-
or 15 weeks after emergence (Menges and Tamez petition of peas. Weeds impaired pea vegetative
1981a). Onion yield did not decrease if the crop was development, especially by reducing tillering in
kept weed free for 2 to 12 weeks after emergence. low-density crops. Therefore, low-density crops had
Climate factors (soil water, soil temperature, and fewer pods per plant at harvest. The presence of the
irradiance) were more useful than weed density to pea crop, independent of its density, did not materi-
explain the differential interference of common sun- ally alter the composition of the weed flora (Lawson
flower in onion between years. and Topham 1985). The crop did not (could not)
Work in irrigated onions in Colorado showed that replace the dominant weed species in high-density
the duration of competition expressed in thermal plots (194 plants m-2 ), but it did reduce the growth
time units with a base of 7.2C explained 65 percent of all species.
of the reduction of onions relative yield (Dunan et Mayweed chamomile produced similar amounts
al. 1996). The first significant reduction in onion of leaf area and dry matter in wet and dry years
yield occurred at 90 thermal time units. A polyno- (Ogg et al. 1993a). Peas, however, produced 20
mial multiple regression model, including duration percent more leaf area and 100 percent more dry
of competition and weed load (weed density and an matter in wet years. The weeds height and dry
estimate of weed competitiveness), accounted for 75 weight increased throughout the growing season,
percent of the variation in relative onion yield. but peas reached a maximum between bloom and
Onions relative yield was more sensitive to the pod set and then declined. Initially, the relative
duration of weed competition than to the specific growth rate of mayweed chamomile was three
weed competitors. times that of pea, but 40 to 48 days after planting
Literature Cited the rates were equal. One might assume early rapid
growth would give the weed an advantage, but the
Dunan, C. M., P. Westra, F. Moore, and P. Chapman.
relative yield of the two species and the relative
1996. Modelling the effect of duration of weed
competition, weed density and weed competitive-
crowding coefficients showed peas were 3 to 20
ness on seeded, irrigated onions. Weed Res. times more competitive and the weed was in fact a
36:359270. weak competitor (Ogg et al. 1993b). In further
Menges, R. M., and S. Tamez. 1981a. Common sun- studies of the same relationship, Ogg et al. (1993a)
flower (Helianthus annuus) interference in onions found root interference was primary and soil water
(Allium cepa). Weed Sci. 29:641747. was more important than nitrogen. If soil water
. 1981b. Response of onion (Allium cepa) to was limiting, mayweed chamomile became more
annual weeds and postemergence herbicides. Weed aggressive than pea. Nitrogen fertilization (20 mg
Sci. 29:7479. wk-1) had no affect on pea yield but more than dou-
bled the weeds size. Pea was the stronger com-
PeaPisum sativum L.
petitor in all cases. The weeds leaf area, root
The sensitivity of beans to weed competition is illus- weight, and shoot weight decreased 55 to 87 per-
trated above in work with redstem filaree (Black- cent with shoot and root interference and 27 to 60
shaw and Harker 1998). Peas were the most percent from only root interference.
82 Chapter 5

Wild mustard competed with a traditional cultivar weeds and vining peas grown at a range of popula-
and a semileafless cultivar that were planted at the tion densities: effects on the crop. Weed Res.
recommended rate of 172 kg ha-1 and at 86 kg ha-1. 23:2728.
Twenty wild mustard m-2 reduced pea seed yield Lawson, H. M., and P. B. Topham. 1985. Competition
nearly the same amount at the two seeding rates (2 between annual weeds and vining peas grown at a
to 35 percent). The semileafless cultivar was more range of population densities. Weed Res.
competitive with the weed at the standard seeding 25:221230.
rate than at the lower rate. Seeding rate had only a ODonovan, J. T., and R. E. Blackshaw. 1997. Effect
modest effect on yield of the traditional cultivar. of volunteer barley (Hordeum vulgare L.) interfer-
ence on field pea (Pisum sativum) yield and prof-
Both cultivars were more affected by the weed in
itability. Weed Sci. 45:249255.
years with normal to high rainfall (Wall et al. 1991).
Ogg, A. G., Jr., R. H. Stephens, and D. R. Gealy.
Volunteer barley reduced pea seed yield 1.7 to 5.4
1993a. Growth analysis of mayweed chamomile
percent over 2 years (ODonovan and Blackshaw (Anthemis cotula) interference in peas (Pisum
1997). This may not seem like a large loss, but it was sativum). Weed Sci. 41:394402.
caused by only 2 to 6 volunteer barley plants m-2. . 1993b. Growth analysis of mayweed
There was no advantage in attempting to manipulate chamomile (Anthemis cotula) and pea (Pisum
pea density above 100 plants m-2 to diminish weed sativum) is affected by form of interference and soil
competition. There was, however, a slight (and water regime. Weed Sci. 42:579585.
unusual among weed-crop studies) economic gain if Wall, D. A., G. H. Friesen, and T. K. Bhati. 1991.
the barley was harvested that could partially offset Wild mustard interference in traditional and semi-
the loss in pea yield. leafless field peas. Can J. Plant Sci. 71:473480.
The beginning of the critical weed-free period in
PepperCapsicum annuum L.
competition with wild oat or tartary buckwheat at
two Canadian locations was 1 or 2 weeks after pea Weed interference periods of 40 to 60 days reduced
emergence (Harker et al. 2001). Weed-free pea bell pepper fruit number and weight 10 and 50 per-
yields at the more northerly location were always cent, respectively (Frank et al. 1992). Foliage weight
two to three times higher than at the second (more of bell pepper declined 10 and 50 percent with inter-
southerly) location. Early competition with tartary ference periods of 20 and 50 days, respectively.
buckwheat at one location in all years did not reduce There was no significant difference in insect infesta-
pea yield, and early competition from wild oat did tion of fruit related to the time of weed interference.
not reduce yield in 1 of 3 years. In general, wild oat Purple nutsedge densities up to 200 plants m-2 lin-
began to reduce pea yield 2 weeks after pea emer- early reduced shoot dry weight at flowering and fruit
gence, and the reduction was linear for the next 2 yield of bell pepper and tomato in Florida as weed
weeks with a loss of 97 kg ha-1 per day. At the sec- density increased (Morales-Payan et al. 1997). For
ond location, early weed competition caused yield each percentage unit of bell pepper shoot dry weight
losses in all years with the onset of losses beginning lost at flowering, fruit yield was reduced 1 to 2.0
1 to 2 weeks after pea emergence. Similar to the first units, with total losses up to 32 percent. Bell pepper
location, yield loss was linear for the next 2 to 3 and tomato both decreased total shoot dry weight of
weeks with a lower average rate of decrease of 45 kg purple nutsedge.
ha-1 per day. Yield losses after full-season competi- The maximum weed infestation period of a natur-
tion ranged from 40 to 70 percent at both sites al weed stand ranged from 0.7 to 3.2 weeks after
(Harker et al. 2001). transplanting to avoid no more than a 5 percent yield
loss in chili pepper in Mexico (Amador-Ramrez
Literature Cited 2002). To prevent total yield decline, weeds had to
Blackshaw, R. E., and K. N. Harker. 1998. Erodium be removed no later than 2.1 weeks after transplant-
cicutarium density and duration of interference ing. However, to prevent a decline of marketable
effects on yield of wheat, oilseed rape, pea, and dry yield, only 0.9 weeks of competition after trans-
bean. Weed Res. 38:5562. planting was permitted. The minimum weed-free
Harker, K. N., R. E. Blackshaw, and G. W. Clayton. period ranged from 6.7 to 15.3 weeks after trans-
2001. Timing weed removal in field pea (Pisum planting with an average of 12.2 weeks of weed-free
sativum). Weed Technol. 15:277283. maintenance to prevent more than a 5 percent yield
Lawson, H. M. 1983. Competition between annual loss.
The Effect of Weed Density 83

Spurred anoda usually emerges in New Mexico Ontario, while the maximum period of weed infes-
after chili peppers are thinned to a final stand tation was 5 to 6 weeks after direct seeding (Weaver
(Schroeder 1993). When 3, 6, 12, 24, or 48 spurred and Tan 1987; Weaver 1984). Thus, the critical peri-
anoda plants were present in 9 m of row, yield od for weed control was between 5 and 9 weeks
decreased 31 to 49 percent when peppers were after direct seeding. A minimum of two weed con-
thinned when they were 10 cm tall and 12 to 27 per- trol operations of some kind were required during
cent when they were thinned when 20 cm tall. the critical period to prevent yield reduction. In con-
Spurred anoda that emerged after thinning trast, Weaver and Tan (1983) demonstrated that the
decreased yield and ease of harvest but not the qual- critical period for weed control in transplanted (as
ity of the harvested crop. opposed to direct-seeded) tomato was 28 to 35 days
Zancada et al. (1998) studied the influence of after transplanting and a single weeding was ade-
root-knot nematode [Meloidogyne incognita quate to prevent yield loss. Yield losses in direct
(Kofoid & White) Chitwood] on competitive inter- seeded tomatoes were attributed to reduction in light
ference between pepper and black nightshade. Very level to tomato by weed shading and weed competi-
few studies of the interaction of other pests and tion for water, which resulted in stomatal closure in
weeds have been done. Root-knot nematode reduced tomato (Weaver and Tan 1987). In transplanted
all growth parameters of pepper but did less harm to tomatoes, growth analysis showed that differences
the weed. Black nightshade was a stronger competi- in plant dry weight and fruit number compared to
tor than pepper with and without nematode infesta- weed-free plots were not apparent until 56 to 70
tion. Nematodes effect on pepper yield was less days after transplanting (Weaver and Tan 1983).
than that of weed competition, but the effects Interference and yield losses were due primarily to
appeared to be additive. shading and not to water stress.
The role of light intensity was affirmed by studies
Literature Cited
of black and eastern black nightshade, which had
Amador-Ramrez, M. D. 2002. Critical period of unequal effects on tomato. Black nightshade was
weed control in transplanted chili pepper. Weed never taller than tomato and did not affect photo-
Res. 42:203209 synthetically active radiation (PAR) at the top of the
Frank, J. R., P. H. Schwartz, Jr., and W. E. Potts. tomato canopy (McGiffen et al. 1992). When densi-
1992. Modeling the effects of weed interference ty of either species increased (up to 4.8 m-2 ), the
periods and insects on bell peppers (Capsicum number of tomato fruits decreased, but eastern black
annuum). Weed Sci. 40:308312. nightshade reduced yield more than black night-
Morales-Payan, J. P., B. M. Santos, W. M. Stall, and shade, because the former was taller and reduced
T. A. Bewick. 1997. Effects of purple nutsedge
PAR at the top of the tomato leaf canopy. PAR at the
(Cyperus rotundus) on tomato (Lycopersicon escu-
top of the tomato canopy was positively correlated
lentum) and bell pepper (Capsicum annuum) vege-
with tomato yield and negatively correlated with
tative growth and fruit yield. Weed Technol.
eastern black nightshade density. Reduction of PAR
Schroder, J. 1993. Late-season interference of spurred during anthesis and early fruit set did not affect
anoda in chile peppers. Weed Sci. 41:172179. tomato yield if PAR during the time of rapid fruit
Zancada, M. C., R. Gonzales-Ponce, and M. Verdugo. development was not reduced (McGiffen et al.
1998. Competition between Solanum nigrum and 1992).
pepper in the presence of Meloidogyne incognita. Losses due to eastern black nightshade and hairy
Weed Res. 38:4753. nightshade always caused greater losses in direct-
seeded than in transplanted tomatoes (Weaver at al.
TomatoLycopersicon esculentum L. 1987). Stomatal conductance and transpiration rates
Work on interference of weeds in tomatoes includes of seeded tomato decreased more rapidly than they
the typical studies that report how many of weed X did in transplanted tomatoes with increasing night-
reduce the yield of tomato by Y amount after Z time. shade density. The value of population density as a
However, work has also been done on competition weed control technique is illustrated by the finding
for light, nutrients, interaction with soil temperature, that seeding tomatoes in twin rows with 33,300 and
and water stress. 45,000 plants ha-1 produced higher yields than those
The minimum weed-free period varied between 7 seeded in single rows with populations of 12,500
and 9 weeks after direct seeding over 3 years in and 22,500 ha-1.
84 Chapter 5

The relative yield of eastern black nightshade and crop growth progressed, tomato shoot dry weight
tomato increased as the proportion of the weed decreased at all barnyardgrass densities. Season
increased in pots, and Perez and Masiunas (1990) long interference decreased fruit number and fruit
concluded that the weed was less competitive than weight at all weed densities (Bhowmik and Reddy
tomato. However, the weed was an effective com- 1988a).
petitor in the field. Tomato yield was reduced by Increasing aggregation of barnyardgrass plants
two-thirds if 3 eastern black nightshade m-1 of row (more clumped distribution) increased intraspecific
grew more than 6 weeks after transplanting. The competition, but interspecific competition from
percentage of marketable tomatoes decreased from tomato decreased (Norris et al. 2001a). The primary
73 without eastern black nightshade to 49 percent influence of different spatial arrangements of the
when the weed was present for 12 weeks. The weed was its effect on shading of tomato. Clumped
importance of weed and crop emergence times was barnyardgrass caused less shading than random or
demonstrated. When eastern black nightshade and regular distribution of the weed along the tomato
tomato were transplanted together, tomato yield was row. With a density of 10 tomatoes m-1 of row, yield
9,000 kg ha-1 and 49 percent of the fruit was mar- losses were 10 to 35 percent in one year and 8 to 50
ketable. If the weed was transplanted 9 weeks after percent in a second year when barnyardgrass was
tomato, yield was 30,000 kg ha-1 and 70 percent of clumped. The same barnyardgrass densities reduced
the fruit was marketable (Perez and Masiunas 1990). tomato yield 20 to 50 percent in one year and 11 to
If tomato was planted late in southeast France, 75 percent in a second year for the regular and ran-
and low densities of black nightshade were present, dom arrangements. Norris et al. (2001a) predicted
two weed-control treatments at the 5- to 6-leaf stage the single-season economic threshold density for a
of crop growth and at the onset of flowering were tomato planting of 10 plants m-1 of row would be 25,
sufficient to prevent yield loss (Caussanel et al. 19, or 15 barnyardgrass plants m-1 of row for the
1990). However, the data also show the vigor of regular, random, and clumped arrangements, respec-
black nightshade competition. If only 12.8 black tively.
nightshade plants ha-1 emerged between the crops 2- When barnyardgrass was grown at densities of 0,
to 3- and 5- to 6-leaf stage, they still caused a sig- 0.25, 0.5, 1, 2, 5, and more than 50 plants m-1 of row
nificant loss if they remained until harvest. in a regular, random, or clumped pattern with toma-
Weaver et al. (1988) compared the relative time of to at densities of 0, 5, 10, or 20 plants m-1 of row in
emergence of four weeds and tomato at five alter- a regular pattern, crop and weed density or spatial
nating temperatures and five levels of available soil arrangement had little effect on phenological devel-
moisture. In general, total emergence decreased for opment of the weed (Norris et al. 2001b). In the
all species as soil moisture decreased, and the absence of tomato, a barnyardgrass plant produced
species differed in the optimum temperature for more than 400,000 seeds without intraspecific com-
emergence, but they were nearly insensitive to soil petition but only 10,000 seeds when plant density
moisture. If one knows the effects of temperature exceeded 50 m-1 of row. Clumped distribution
and moisture on weed species, this information can reduced seed production 30 to 50 percent when den-
be used to develop optimal crop planting dates and sity was 1 to 5 plants m-1 of row. Tomato interference
to estimate potential crop yield losses (Weaver et al. reduced barnyardgrass seed production, but the
1988). magnitude of reduction depended on tomato and
Prevailing weather influenced the competitive weed density. Nearly 700,000 seeds m-2 were pro-
effect of jimsonweed, tall morningglory, and com- duced when the weeds density exceeded 50 plants
mon cocklebur about equally (Monaco et al. 1981). m-1 of row. The most significant conclusion of these
If they were present for the whole season, densities studies (Norris et al. 2001a, b) was that barnyard-
of 11, 43, or 86 plants m-2 reduced tomato yield in a grass seed production at the single-season economic
warm year. The second year of study was wetter, threshold density was sufficient to maintain the soil
slightly cooler, and the crop was irrigated. Then den- seedbank and require high levels of weed control in
sities between 2.7 and 11 of each weed m-2 reduced subsequent years. Preventing seed production was
yield. recommended as the best long-term management
Barnyardgrass density of 16 plants m-1 of row strategy, a recommendation that is biologically and
reduced tomato yield 26 percent, while a density of economically wise but one that has not been widely
64 weeds m-1 of row reduced yield 84 percent. As accepted.
The Effect of Weed Density 85

When common lambsquarters and wild mustard between the years. Crop growth rate and above-
were grown with tomato in additive and replacement ground dry biomass of tomato cultivars grown with
series studies in the greenhouse, both species had velvetleaf were generally less than when the culti-
similar effects on shoot dry weight at low densities (2 vars were grown in monoculture. Yield loss with
or 5 plants per pot) but wild mustard was more com- high weed density was similar among the four culti-
petitive at higher densities (15 to 20 plants per pot) vars but it was variable at low weed density.
(Quasem and Hill 1994). Weed competition did not Relative yield analysis indicated that tomato is a
affect N, P, K, Ca, or Mg content of tomato shoots, stronger competitor than either purple or yellow
but total tomato dry matter and total nutrients were nutsedge. Both nutsedge species are strong intraspe-
reduced with increasing density of both weeds. These cific competitors (Santos et al. 1997). When either
findings are similar to the earlier study of Sanders et nutsedge species was grown with tomato for 40 days,
al. (1981), which found few instances of differences tomato dry weight per plant increased and that of the
in nutrient content of tomato and the leaf tissue of nutsedges decreased. This was due primarily to the
jimsonweed, tall morningglory, common cocklebur, ability of the tomato leaf canopy to shade nutsedge,
or large crabgrass. There was no clear relationship which is particularly susceptible to shading. Field
between the concentration of N, P, K, Ca, Mg, or S experiments demonstrated that the fungus Dactylaria
and weed density. More tomato fruit was produced in higginsii, a native of Florida, isolated from purple
weed-free plots per kg of total assimilated N, P, and nutsedge, produced tomato yields equivalent to a
K than in weedy plots (Sanders et al. 1981). Quasem weed-free control when 106 conidia ml-1 were applied.
and Hill (1994) reported N, P, K, and Mg concentra- Purple nutsedge was present in the pots at densities of
tions were higher in shoots of common lambsquar- 40, 80, 160, or 320 tubers m-2 (Kadir et al. 1999).
ters. Reduction of growth of common lambsquarters Another of the very few studies of interference
was associated with a reduced ability to accumulate from parasitic weeds investigated the effect of les-
K. The competitive index of common lambsquarters pedeza dodder on tomato (Goldwasser et al. 2001).
decreased with its proportion in a mixture but the The primary finding was that some tomato commer-
opposite was true for wild mustard. Common lambs- cial hybrid varieties are at least partially resistant or
quarters was nearly 3.5 times as competitive as wild tolerant to parasitism by dodder. In the field, dodder
mustard, and their relative competitive ability was seed germinated, emerged, and began to twine
closely related to the growth of their root systems, a around tomato stems, but in tolerant cultivars, haus-
factor overlooked in many competition studies. toria attachments were 75 percent less and dodder
In another study, common lambsquarterss densi- growth was reduced up to 70 percent.
ty ranged from 16 to 64 plants m-1 of row, and the
Literature Cited
weed fresh weight ranged from 26,360 kg ha-1 with
16 plants ha-1, to 46,000 kg ha-1 with 64 weeds m-1 of Bhowmik, P. C., and K. N. Reddy. 1988a. Effects of
row. Season-long interference of common lambs- barnyardgrass (Echinochloa crus-galli) on growth,
quarters varied from 17 percent at the low density to yield, and nutrient status of transplanted tomato
36 percent yield loss at the high density. Nitrogen (Lycopersicon esculentum). Weed Sci. 36:775778.
concentration in tomato leaves was unchanged dur- 1988b. Interference of common lambsquarters
ing the vegetative and flowering stages but declined (Chenopodium album) in transplanted tomato
regardless of the weeds density at the early-fruit (Lycopersicon esculentum). Weed Technol.
stage and at harvest (Bhowmik and Reddy 1988b). 2:505508.
Similar to other studies, weed density did not affect Caussanel, J. P., X. Branthome, J. Maillet, and A.
Carterton. 1990. Influence de la densit et de la
P, K, or Ca levels in tomato leaves.
priode de concurrence de Solanum nigrum L. sur
Field experiments in California were conducted
la tomate de semis direct, en relation avec le
with four tomato cultivars to determine if there were
dsherbage. Weed Res. 30:341354.
cultivar traits that could be associated with compet- Goldwasser, Y., W. T. Lanini, and R. L. Wrobel. 2001
itiveness against velvetleaf (Ngouaijo et al. 2001). tolerance of tomato varieties to lespedeza dodder.
The weeds competitive effects varied with year and Weed Sci. 49:520523.
cultivar. When velvetleaf density was 5 m-1 of row, Kadir, J. B., R. Charudattan, W. M. Stall, and T. E.
the yield of one cultivar was reduced 8 percent in Bewick. 1999. Effect of Dactylaria higginsii on
one year and 60 percent in a second year. For anoth- interference of Cyperus rotundus with L. Esculen-
er cultivar, the variation was 58 to 80 percent tum. Weed Sci. 47:682686.
86 Chapter 5

McGiffen, M. E., Jr. , J. B. Masiunas, and J. D. Hes- Other Vegetable Crops

keth. 1992. Competition for light between tomatoes
and nightshades (Solanum nigrum or S. CabbageBrassica oleracea L.
ptycanthum). Weed Sci. 40:220226. Cabbage yield was reduced if plots were not kept
Monaco, T. J., A. S. Grayson, and D. C. Sanders. weed free for at least 3 weeks after planting or if
1981. Influence of four weed species on the growth, weeds competed more than 4 to 5 weeks after plant-
yield, and quality of direct-seeded tomatoes (Lycop- ing (Weaver 1984). If one attempted to manage
ersicon esculentum). Weed Sci. 29:394397. weeds by planting a higher crop population (nar-
Ngouaijo, M., M. E. McGiffen, Jr., and K. J. Hem- rower rows), it was counterproductive because the
bree. 2001. Tolerance of tomato cultivars to vel-
result was smaller crop plants and earlier competi-
vetleaf interference. Weed Sci. 49:9198.
tion from weeds and a shorter time during which the
Norris, R. F., C. L. Elmore, M. Rejmnek, and W. C.
crop could withstand weed competition. Weaver
Akey. 2001a. Spatial arrangement, density, and
competition between barnyardgrass and tomato. I.
(1984) did not identify a true critical period for
Crop growth and yield. Weed Sci. 49:6168. weed competition in cabbage.
. 2001b. Spatial arrangement, density, and Miller and Hopen (1991), in Wisconsin, identified
competition between barnyardgrass and tomato. II. the critical weed-control period in cabbage as 2
Barnyardgrass growth and seed production. Weed weeks in one year and 4 weeks in a second year with
Sci. 49:6976. a natural weed stand. Season-long velvetleaf densi-
Perez, F. G. M., and J. B. Masiunas. 1990. Eastern ties of 1.2 or 3.6 plants m-2 reduced cabbage yield 52
black nightshade (Solanum ptycanthum) interfer- and 76 percent in one year and 76 to 92 percent in a
ence in processing tomato (Lycopersicon esculen- second year. All velvetleaf densities planted 0, 1, or
tum). Weed Sci. 38:385388. 2 weeks after cabbage reduced cabbage yield, but
Quasem, J. R., and T. A. Hill. 1994. Inter- and planting 4 or 6 weeks after cabbage had no effect on
intraspecific competition of fat-hen (Chenopodium yield.
album) and groundsel (Senecio vulgaris L.). Weed A study in the subtropical environment of
Res. 34:109118. Taichung, Taipei, determined the effect of planting
Sanders, D. C., A. S. Grayson, and R. F. Mumm. cabbage in fields after rice harvest (Horng 1980) to
1981. Mineral content of tomato (Lycopersicon give cabbage a longer growing period. Cabbage was
esculentum) and four competing weed species. transplanted into rice fields 3 to 12 days before rice
Weed Sci. 29:590593. harvest or 5 days after harvest. Cabbage transplant-
Santos, B. M., T. A. Bewick, W. M. Stall, and D. G. ed before rice harvest began to grow before pale
Shilling. 1997. Competitive interactions of tomato
smartweed had germinated. Its yield was reduced
(Lycopersicon esculentum) and nutsedges (Cyperus
when the weed was allowed to grow more than 4
spp.). Weed Sci. 45:229233.
weeks from the time beds were formed around cab-
Weaver, S. E. 1984. Critical period of weed competi-
tion in three vegetable crops in relation to manage-
bage which was about 5 days after rice harvest. If
ment practices. Weed Res. 24:317326. cabbage was transplanted 5 days after rice harvest,
Weaver, S. E., N. Smits, and C. S. Tan. 1987. Estimat- pale smartweed had emerged and cabbage yield was
ing yield losses of tomatoes (Lycopersicon esculen- reduced when the weed grew for only 2 weeks after
tum) caused by nightshade (Solanum spp.) beds were formed. Cabbage has to be kept weed free
interference. Weed Sci. 35:163168. for a minimum of 4 weeks after transplanting to pre-
Weaver, S. E., and C. S. Tan. 1983. Critical period of vent yield reduction (Horng 1980).
weed interference in transplanted tomatoes (Lycop-
ersicon esculentum): Growth analysis. Weed Sci. CucumberCucumis sativus L.
. 1987. Critical period of weed interference A natural infestation of yellow nutsedge with densi-
in field-seeded tomatoes and its relation to water ties up to 955 plants m-2 had a 5 percent reduction in
stress and shading. Can J. Plant Sci. cucumber yield with a yellow nutsedge density of
67:573581. about 15 plants m-2. Yellow nutsedge was a more
Weaver, S. E., C. S. Tan, and P. Brain. 1988. Effect of effective competitor in uneven or nonuniform
temperature and soil moisture on time of emer- cucumber stands. Uniform cucumber stands maxi-
gence of tomatoes and four weed species. Can J. mized the crops competitive ability (Johnson and
Plant Sci. 68:877886. Mullinix 1999).
The Effect of Weed Density 87

Cucumber yield was reduced if plots were not when competition was intraspecific and four times
kept weed free for up to 4 weeks after planting or if more when it was interspecific. Spiny amaranth was
weeds competed more than 3 to 4 weeks after plant- more competitive than lettuce regardless of the
ing (Weaver 1984). If one attempted to manage phosphorus level, but phosphorus increased the
weeds by planting a higher crop population (nar- competitiveness of lettuce. The total lettuce shoot
rower rows), it was counterproductive because the biomass per pot and the weight per plant increased
result was smaller crop plants and earlier competi- 39 and 44 percent in response to phosphorus (Shre-
tion from weeds and a shorter time during which the fler et al. 1994a).
crop could withstand weed competition. Weaver
(1984) did not identify a true critical period for RadishRaphanus sativus L.
weed competition in cucumbers. Radish and shepherds-purse have similar height,
LeekAllium ampeloprasum L. leaf area, and root biomass, and they often occur
together. In mixtures, radish was the stronger com-
Leek and most Allium spp. (e.g., onion, garlic) are petitor. Its total dry matter and tuber production
weak competitors. They have minimal shoot struc- were affected only slightly or not at all in mixtures
ture and do not readily cover the soil surface. Stud- with shepherds-purse. In monoculture, the two
ies were done to determine if celery could be used as species had similar leaf areas, but that of shepherds-
a companion crop to suppress weeds in leek (Bau- purse was greatly reduced in mixtures because of
mann et al. 2000). Intercropping leek and celery in a the ability of radish to intercept light. Radish was
row-by-row design shortened the critical period for able to grow taller in mixture than it did in mono-
weed control in the intercrop compared to culture (Perera and Ayres 1992).
monocropped leek. The relative soil cover of weeds
that had emerged at the end of the critical period was Literature Cited
reduced by 41 percent in the intercrop. The biomass Baumann, D. T, M. J. Kropff, and L. Bastiaans. 2000.
of common groundsel that was planted 20 days after Intercropping leeks to suppress weeds. Weed Res.
crop establishment was reduced 58 percent in the 40:359374.
intercrop and seedlings emerging from the planted Horng, Liang-Chi. 1980. Interference of pale
common groundsel were reduced 98 percent com- smartweed (Polygonum lapathifolium) with cab-
pared to monocropped leek. In addition, the relative bage (Brassica oleracea). Weed Sci.
yield total of the intercrop exceeded the pure stand 28:381384.
by 10 percent, which Baumann et al. (2000) attrib- Johnson, W. C., III, and B. Mullinix, Jr. 1999. Cyper-
uted to more optimal use of resources. However, us esculentus interference in Cucumis sativus. Weed
leek quality was reduced. The idea of intercropping Sci. 47:327331.
or companion cropping for weed management is not Miller, A. B., and H. J. Hopen. 1991. Critical weed-
new but its potential has not been adequately inves- control period in seeded cabbage (Brassica oler-
tigated. acea var. capitata) Weed Technol. 5:852857.
Perera, K. K., and P. G. Ayres. 1992. Effects of shep-
LettuceLactuca sativum L. herds purse [Capsella bursa-pastoris (L.) Medic]
Seven weeks of interference from spiny amaranth on the growth of radish (Raphanus sativus L.).
reduced lettuce head weight 20 percent and 8 weeks Weed Res. 329336.
Shrefler, J. W., D. G. Shilling, J. A. Dusky, and B. J.
reduced head weight 24 percent. When phosphorus
Brecke. 1994a. Influence of phosphorus fertility on
was banded, the effect of spiny amaranth was
intra- and interspecific interference between lettuce
reduced, but the interference between the species
(Lactuca sativa) and spiny amaranth (Amaranthus
was not due to competition for phosphorus. The spinosus). Weed Sci. 42:574578.
weeds density and the duration of interference had Shrefler, J. W., J. A. Dusky, D. G. Shilling, B. J.
little to no effect on the phosphorus content of let- Brecke, and C. A. Sanchez. 1994b. Effects of phos-
tuce (Shrefler et al. 1994b). The addition of phos- phorus fertility on competition between lettuce
phorus made lettuce and spiny amaranth equally (Lactuca sativa) and spiny amaranth (Amaranthus
competitive at low densities but spiny amaranth was spinosus). Weed Sci. 42:556560.
four times more competitive at high densities in a Weaver, S. E. 1984. Critical period of weed competi-
greenhouse/pot study (Shrefler et al. 1994a). The tion in three vegetable crops in relation to manage-
weed produced 2.4 times more biomass than lettuce ment practices. Weed Res. 24:317326.
88 Chapter 5

WHEATTRITICUM AESTIVUM L. parable or greater in no-till plots and wheat produc-

tion was maintained in spite of reduced plant estab-
This chapter claimed (see Rice, above) that rice is lishment (Lgre and Bai 1999).
the worlds most important crop because more peo- Cousens and Mokhtari (1998) studied the ability
ple depend on it for their daily sustenance than on of wheat cultivars to maintain yield in the presence
any other crop. If the number of acres on which a of weeds over several locations and in successive
crop is grown or the range of latitude over which the years. They found that the magnitude of the yield
crop is grown are the appropriate criteria, then advantage for some cultivars differed a lot between
wheat, not rice, is the worlds most important crop. years and locations. More important, there was little
It is grown on about 213.6 million ha each year over correlation between competitiveness at different
a wider latitudinal range than other major crops. It is sites in one year or between years. Cousens and
grown where the weather is too dry or too cold for Mokhtari (1998) found only one cultivar that was a
rice or corn. The annual crop is 576.3 million metric consistently good competitor and two that were con-
tons with an average worldwide yield of 2,698 kg sistently poor. Their quest was to identify cultivars
ha-1 (United Nations 2000). Clearly wheat is one of that were consistently more competitive so they
the worlds most important crops. could be recommended to farmers, but their data did
Challaiah et al. (1983, 1986) used a field study to not provide any basis for consistent, good advice to
select wheat cultivars that were competitive with farmers.
downy brome. Downy brome reduced the grain Wicks et al. (1994) suggested that winter wheat
yield of all cultivars 9 to 21 percent at one site and cultivars that are more competitive help control
20 to 41 percent at a second site in the same year. weeds in rotational crops. Cultivars that averaged 90
Wheat tiller number, canopy diameter and height to 109 cm tall were consistently more competitive
were all negatively correlated with downy brome than those that were 80 to 89 cm or 69 to 79 cm tall.
yield, but changes in these parameters did not When sorghum was grown in fields that had grown
always result in an increase in wheat grain yield. In the more competitive wheat cultivar, weed biomass
this study (Challaiah et al. 1983, 1986), wheat in sorghum was 61 percent less than in fields that
height was most closely correlated with a decrease had grown the less-competitive wheat cultivars.
in downy brome yield. Sorghum yield was also higher when it was grown
Lgre and Bai (1999) included rapid and uni- after more-competitive wheat cultivars.
form seedling emergence, tillering, early biomass Christensen (1994) used oilseed rape as a substi-
accumulation, canopy closure, and a height advan- tute for weeds to determine if there was a significant
tage over competing weeds to evaluate the effects of interaction between the competitive ability of wheat,
no-tillage practices on growth and productivity of barley and rye cultivars and herbicide performance.
oats, barley, and wheat. All three crops were grown A target level of 5 g of weed dry matter m-2 was
with and without soil tillage, and the cereal growth used. He found that one winter wheat cultivar
parameters were measured six or seven times during required a 154 percent higher herbicide dose than a
the growing season. Grain yield and yield compo- winter barley cultivar, whereas for winter rye, herbi-
nents were determined at crop maturity. Oats and cide dose could be reduced 31 percent.
barley were little affected by tillage but wheat pop- Rather than crop cultivars, Wilson and Wright
ulations were reduced 16 to 20 percent by no-tillage. (1990) studied the role of growth and competitive-
Height in no-tilled systems was similar or greater ness and ranked the competitive order of 12 annual
than in tilled systems for all three cereals (Lgre weeds in winter wheat. Weeds that senesced in mid-
and Bai 1999). Leaf area index and biomass accu- summer were less competitive than those with a
mulation were also comparable in both systems for growth pattern similar to wheat. In one year, most
all three cereals, except for wheat, which was weeds had little effect on wheat because crop densi-
greater in tilled systems but only on two sampling ty was high. Crop yieldweed density relationships
dates. Lgre and Bai (1999) found that the response for all species in one year and for catchweed bed-
of annual dicot weeds to tillage was inconsistent in straw in a second year were described well by the
all crops. Perennial dicots dominated no-tillage sys- rectangular hyperbola. Wilson and Wright (1990)
tems and perennial monocots were more abundant proposed that a competitive index, derived from
in tilled systems for all three cereals. Yield of all yield density relationships and expressed as the per-
three cereals (except barley in one year) was com- cent yield loss for a weed m-2, is more likely to
The Effect of Weed Density 89

reflect a weeds competitive ability in a crop than is wheat and sterile oat. Dry weight of wheat, barley,
an index derived from a plants weight in the crop. and triticale were not affected by sterile oat (110
Some of these relationships may be explained by plants m-2) until March of the year after planting, but
the work of Barnes et al. (1990), who demonstrated yields were reduced by sterile oat competition after
the importance of canopy structure as a determinant that time. Grain yield of wheat and barley were
of the light interception and carbon gain in mixed reduced 61 percent by 110 sterile oats m-2 but barley
and pure stands of wheat and wild oat. In mixtures, yield was reduced only 9 percent. Nitrogen (150 kg
the fraction of the simulated canopy photosynthesis ha-1) slightly increased yield of all crops in mono-
contributed by wheat declined during the growing culture, and it increased sterile oats dry weight and
season and the decline was closely related to reduc- its competitive ability against wheat and triticale
tion in the amount of leaf area in the upper canopy. (Dhima and Eleftherohorinos 2001). When 50 kg of
Canopy photosynthesis for both species was most nitrogen ha-1 was applied before planting and fol-
sensitive to change in leaf area and leaf inclination lowed by 100 kg ha-1 in early March, there was a
in the middle and upper canopy. Changes in leaf slightly higher increase in sterile oats dry weight
area index and leaf inclination affected carbon gain compared to one nitrogen application before plant-
differently in mixtures and monoculture and differ- ing. The results indicate that, for winter wheat in
ently for the two species (Barnes et al. 1990). Total Greece, the time of nitrogen application could be
leaf area alive and functioning at one time in each used to give a slight advantage to the crop and that
species was the most important determinant of com- barley is more effective at limiting a sterile oat infes-
petitive success. Light competition in the mixture of tation than wheat or triticale.
wheat and wild oat was influenced heavily by dif- The competition between sterile oat and six wheat
ferences in positioning of leaf area in the upper cultivars, each with a different maturity time, was
canopy, which determines the amount of light inter- studied in a greenhouse experiment by Gonzlez-
cepted (Barnes et al. 1990). Ponce (1988). He determined that competitiveness
Wells (1979) studied the effect of the density of was similar for all cultivars, but the cultivars with
five broadleaved weed species at five different sites the longest time to maturity were affected most
on the yield of wheat at three levels of applied nitro- because they were consistently shorter. This is
gen. The relationship between dry matter production because most of the stem extension and some of the
and population density for all but one of the five grain formation took place after the sterile oat pani-
weeds was curvilinear but the curvature was slight cles had expanded above the wheat canopy.
and the effects of competition of the weeds in wheat Intraspecific competition was always greater than
was linear for four of the five weeds. Nitrogen interspecific competition.
increased wheat yield at all sites but the effects of The interference of four weeds in wheat has been
weed competition did not change (Wells 1979). studied more than all of the 25 other weeds that have
Gill and Blacklow (1984) investigated the effect been studied. The four weeds are the ryegrasses
of great brome on the growth of wheat and its uptake (especially Italian ryegrass), Bromus spp. (especial-
of nitrogen and phosphorus. Shoot dry matter of ly downy brome), blackgrass, and jointed goatgrass.
wheat was reduced from 1.4 g per plant to 0.5 g per None of these has been studied as extensively as
plant after it competed for 71 days with 400 great other species have been studied in corn and soybean.
brome m-2. Competition with 400 great brome m-2 Liebl and Worsham (1987) studied the interfer-
reduced the concentration of nitrogen in wheat ence of Italian ryegrass in wheat in North Carolina.
shoots with three tillers (Feekes scale 3) from 4.1 to Wheat yield declined 4.2 percent for every 10 Ital-
3.2 g and phosphorus from 0.77 to 0.58 g. Gill and ian ryegrass plants m-2 when the weed density
Blacklow (1984) determined that nitrogen and phos- ranged from 0 to 100 m-2. The weeds effect was pri-
phorus concentrations in wheat shoots were reduced marily a reduction in wheat tillering. Italian ryegrass
before any detectable reductions in dry matter. One densities as high as 80 m-2 had little to no effect on
can conclude that great brome competed with wheat wheat head or kernel weights. Liebl and Worsham
for these nutrients, but competition for water during (1987) also found that Italian ryegrass responded
grain filling caused the greatest reduction in grain more than wheat to added nitrogen and potassium.
yield. The net uptake of the two nutrients was twice as
Dhima and Eleftherohorinos (2001) studied the great for the weed as it was for wheat, so there was
influence of nitrogen on competition between winter probably some level of competition for nutrients.
90 Chapter 5

Italian ryegrass had significantly greater biomass in during wheats reproductive stages. This was
monoculture than in competition with wheat because 200 days after emergence the leaf area
because wheat seedlings were much larger than the index of Italian ryegrass was 6.6 times that of wheat.
weed seedlings for the first 20 days after emergence. The weed reduced the photosynthetically active
Thus, the results of this study were affected by the radiation reaching wheat up to 68 percent at wheats
weeds effect on tillering of wheat, some competi- booting stage (Hashem et al. 1998). Wheats grain
tion for nutrients, and wheats initial advantage of yield could be reduced up to 92 percent by Italian
greater seedling size. ryegrass. As few as 9 weeds in 100 wheat plants
Ghersa et al. (1994) (in Argentina and Oregon) m-2 reduced wheat yield as much as 33 percent. In a
took a different approach and asked if the radiation subsequent study, Hashem et al. (2000) looked at the
environment during winter-wheat establishment competitive effect of winter wheat planted in a
could be manipulated to favor wheat. The percent- square arrangement with Italian ryegrass planted
age of total radiation and the ratio of red (660 nm) randomly, on biomass yield of each species, Italian
to far-red light (730 nm) that reached the soil surface ryegrass seed yield, nitrogen use efficiency, and
were important regulators of Italian ryegrass germi- progeny seed germination. Increases in wheat densi-
nation, growth, and subsequent interference with ty up to 800 plants m-2 reduced Italian ryegrass seed
wheat. If the total radiation that reached the soil sur- yield up to 87 percent but increased its harvest index
face was reduced to about 10 percent of full sunlight up to 42 percent. Density and species interaction
while the red:far red ratio was maintained at about accounted for 66 to 73 percent of the total variation
1.0, wheat grain yield fell 40 percent compared to in each species biomass, and association (Italian
weed-free wheat in full sunlight. Reducing total ryegrass and wheat growing together) was less
radiation to only 3 percent of full sun and reducing harmful to Italian ryegrass. Both inter- and intraspe-
the red:far red ratio to 0.2 reduced wheat grain yield cific competition increased nitrogen content of
in the presence of Italian ryegrass by 35 percent wheat grain, whereas only interspecific competition
compared to wheat yield in full sun. In both the pure affected nitrogen content of Italian ryegrass seed.
Italian ryegrass and the mixed stands, Italian rye- Hashem et al. (2000) found that wheats nitrogen
grass dry matter production was reduced by low uptake was three times greater than that of Italian
irradiance and the low red:far red ratio. The combi- ryegrass, but the latter was twice as efficient as
nation of low irradiance and low red:far red ratio wheat was at producing a unit of biomass per unit of
reduced dry matter production to about 50 percent nitrogen. In contrast to the results of Dhima and
of that in the control. Red light enrichment beneath Eleftherohorinos (2001), nitrogen was not the main
the plant canopy, to achieve a red:far red ratio of 1.3, limiting factor in competition between the two
increased Italian ryegrass germination by 20 percent species.
compared to normal light. Ghersa et al. (1994) did Stone et al. (1998) examined aboveground and
not define the mechanism, but clearly demonstrated below-ground interference of Italian ryegrass and
that manipulation of radiation during the early wheat in the greenhouse. They found that above-
stages of crop growth may be a good weed manage- ground interference of Italian ryegrass had no effect
ment technique. on wheat and that the volume of soil (pot volume of
Hashem et al. (1998) used an addition series 90, 950, or 3,800 ml) in which 1 wheat plant grew
experiment to study interference of Italian ryegrass with 9 Italian ryegrass plants had no effect on inter-
in wheat. In monoculture, 80 to 85 percent of the ference interactions. However, if Italian ryegrass
variation in wheat biomass was explained by crop interference was restricted to roots, the weed had a
density (plants m-2 ). However, in mixtures, initial competitive advantage.
wheat density and initial weed density explained 74 Carson et al. (1999) considered the relative abili-
to 80 percent of wheat total biomass and 68 to 79 ty of wheat and Italian ryegrass in adequately
percent of the total biomass of Italian ryegrass. watered and dry environments. When Italian rye-
Interspecific competition was apparent 15 to 90 days grass was grown alone for 14 weeks, it produced a
after emergence for wheat but not until 90 to 170 greater leaf area, greater dry weight of stem and
days after emergence for Italian ryegrass. The max- roots, and more tillers than monocultural wheat.
imum intraspecific competition occurred in wheat Because wheat had larger seeds, grew taller, and had
170 days after emergence, whereas maximum inter- a larger initial leaf area, it was able to produce a
specific competition with Italian ryegrass occurred greater final leaf area and stem dry weight when
The Effect of Weed Density 91

grown with Italian ryegrass. Wheat was able to the three major nutrients and for light. Rigid rye-
maintain a greater leaf expansion rate during grass dry matter and seed production were negative-
drought and a greater leaf area afterward and thus ly correlated with yield of each crop. Lemerle et al.
had greater growth than Italian ryegrass in pure cul- (1995) concluded that competitive crops offered
tures of each. But when drought was relieved, the promise for suppression of grass weeds, especially
relative competitive ability of Italian ryegrass in in the case of grain legumes.
mixtures with wheat was enhanced. Medd et al. (1985) determined that wheats plant-
In contrast to other studies, Tanji et al. (1997) ing arrangement had little influence on rigid rye-
showed in greenhouse studies and separate field grass competition independent of crop density.
studies in Morocco that wheat was an effective com- However, rigid ryegrasss effect on wheat yield was
petitor with Italian ryegrass and cow cockle. One substantially reduced by increasing wheat plant den-
wheat plant was as competitive as 11 Italian rye- sity from 40 or 75 to 200 plants m-2. The reciprocal
grass plants in the greenhouse and as 19 Italian rye- yield model was a good predictor of yield reduction,
grass in the field. One wheat plant was as especially when it included the ratio of weed to crop
competitive as 3 or 24 cow cockle plants. Shoot dry density. Lemerle et al. (2001) also studied the rela-
weight was the easiest, fastest, and least-expensive tive competitive advantage of 12 commercial wheat
parameter to measure, and it was employed, but, as varieties at several sites in southeastern Australia
other studies have shown (e.g., Hashem et al. 1998; over 3 years (see Cousens and Mokhtari 1998 and
Stone et al. 1998), it may not have been the best Christensen 1994 for similar work). Nearly all (81
parameter to measure. Growth analysis (Tanji et al. percent) the variation in wheat yield was attributed
1997) showed, as Carson et al. (1999) showed, that to variety x environment effects with only 4 percent
wheat had a greater leaf area, shoot and root dry due to variety x weed x environmental effects. Three
weight, and absolute growth rate than either weed, varieties exhibited environment-specific competitive
especially early in the growing season. Tanji et al. advantages, and at least three others were poor com-
(1997) concluded that if wheat density was 120 to petitors in some environments. Lemerle et al. (2001)
240 m-2, wheat alone could minimize the competi- proposed that greater genetic variability was
tion of each weed and acceptable wheat yields could required in wheat to improve competitiveness. They
be obtained under Moroccos dryland conditions. also suggested the older tactic of increasing wheat
Two papers have explored interference of two seeding rate as a useful management technique.
other ryegrass species: rigid ryegrass (Lemerle et al. Varieties that showed a competitive yield advantage
2001) and perennial ryegrass (Wright and Hebbleth- also suppressed growth of rigid ryegrass. Lemerle et
waite 1983). The paucity of papers on rigid ryegrass al. (2001) concluded that Australian wheat breeders
was surprising given its importance in Australia. may inadvertently select for competitive advantage
However, much of that work has emphasized control against weeds when they select for traits such as
and management in light of problems with herbicide early vigor.
resistance and that work is not reviewed here. Wright and Hebblethwaite (1983) demonstrated,
Lemerle et al. (1995) studied the competitive abil- as others have, that perennial ryegrass seed yield in
ity of eight winter crops with the annual, rigid rye- the presence of wheat depended on wheat density
grass. The purpose was to determine if one or more and the number and weight of perennial ryegrass
of the crops could be used as part of a weed man- tillers (its density). The greatest reduction in peren-
agement program to suppress rigid ryegrass. The nial ryegrass seed yield occurred, as expected, in the
order of decreasing competitive ability with the presence of high (300 plants m-2) wheat density.
range of percent yield reduction for each species in When wheat density was low (80 plants m-2), rye-
competition with 300 rigid ryegrass plants m-2 was: grass seed yield was reduced much less, but the
oats (214 percent), cereal rye (1420 percent), number of live wheat plants gradually decreased.
oilseed rape (930 percent), wheat (2240 percent), Between 0 and 300 perennial ryegrass m-2, a 1 per-
barley (1055 percent), field pea and lupine (100 cent loss of perennial ryegrass seed yield occurred
percent). There were differences in the competitive for every 10 wheat plants m-2. The work demon-
ability of the two cultivars chosen for each species, strates the effectiveness of crop density as a weed
but competition was more strongly influenced by the management technique.
different growing conditions in each year. The study Blackgrass has been a much greater concern in
(Lemerle et al. 1995) demonstrated competition for the UK than in the United States. The seriousness of
92 Chapter 5

the management problem was illustrated by Moss emergence. Downy brome that emerged 21 or more
(1983) who showed that over 95 percent of black- days after wheat did not affect wheat yield. With
grass seed was shed before winter wheat was har- downy brome densities up to 100 m-2, the quadratic
vested, and 70 percent of all seed produced was shed equation best described wheat yield loss as a func-
between mid-July and mid-August prior to wheat tion of weed density when the weed emerged within
harvest. Barley, because it is harvested a bit earlier 14 days of wheat.
in the UK, had fewer seeds shed prior to harvest. Blackshaw (1993) affirmed that the time of
Wilson (1979) stressed the need for early control, downy brome emergence relative to wheat emer-
especially when blackgrass and wild oats were both gence affected the magnitude of wheat yield reduc-
present in wheat. tion, and time of emergence was more important
In northern Turkey, Mennan et al. (2002) deter- than downy brome density. Blackshaw (1993)
mined that the economic threshold for blackgrass showed that when wheat and downy brome had
was 23 to 39 plants m-2, whereas it was only 11 to 20 comparable density, the weed caused two- to five-
plants m-2 for wild oat, at densities of 2, 5, 10, 20, or fold greater reductions in yield when it emerged
40 plants m-2. within 3 weeks after wheat than when it emerged 6
The effect of weed density, nitrogen fertilizer, weeks after wheat or in early spring. Late-emerging
crop planting date, and weed emergence on compe- downy brome could cause significant wheat yield or
tition between wheat and blackgrass was studied in biomass losses, but only at densities of 200 to 400
the greenhouse by Exley and Snaydon (1992). Root plants m-2. The lack of effect of late-emerging
competition affected the growth of wheat and black- downy brome was due to shading (70 to 90 percent
grass more than shoot competition, although shoot light reduction) of the weed by wheat for most of the
competition affected blackgrass survival more than growing season. When downy brome emerged early,
root competition. Nitrogen fertilizer partially allevi- it could reduce wheat biomass up to 59 percent and
ated the effects of root competition but did not affect wheat seed yield up to 68 percent (Blackshaw
root competition because each species has similar 1993). In a subsequent study, Blackshaw (1994)
needs. If either species emerged later than the other, clearly demonstrated that crop rotation and soil
its competitive ability was reduced, particularly management influence downy brome density. In
when competition was restricted to roots. continuous wheat, downy brome density increased
Work by Melander (1995) did not quite confirm from 24 to 970 plants m-2 over 5 years. Weed densi-
the effect of planting date on competitive ability of ties were often higher with no-tillage. When fallow
blackgrass and silky bentgrass. Melander used two or spring canola were used in rotation with winter
planting dates spaced 14 to 16 days apart. Planting wheat, downy brome density decreased to less than
date had an inconsistent effect on weed populations 55 to 100 plants m-2, respectively, over 6 years.
in spring, but seedling emergence of both weeds Clearly, as Blackshaw points out (1994), continuous
appeared to be delayed when later and early planting winter wheat will only worsen the downy brome
were compared. The delay could partially explain problem in areas where the weed is prevalent, and,
the reduced competitive ability of both weeds with therefore, crop rotation is a desirable management
the later planting (emergence) date. In some cases, practice.
seed population per plant was lower with later plant- Koscelny et al. studied the effects of winter wheat
ing, but in other cases, there was no effect. Reduced row spacing, cultivar, seeding rate, water, and nitro-
seed production by each weed was caused more by gen on competitiveness of hard red winter wheat
a reduction in the number of inflorescences per plant with cheat (1990). In a separate experiment, they
than by planting date. In spite of variable results, examined the interaction of wheat seeding rate and
Melander (1995) concluded that planting dates row spacing on interference by cheat (1991). When
should be considered as a useful part of any man- row spacing decreased from 23 to 8 cm, yield of
agement program for annual grass weeds in small weed-free wheat increased at two of three locations
grains. and the yield of cheat-infested wheat increased in 6
The interference of three Bromus species has been of 10 experiments. Increasing seeding rate from 265
studied in winter wheat. Stahlman and Miller (1990) to 530 seeds m-2 increased wheat yield. No wheat
found that densities of 24, 40, or 65 downy brome cultivar consistently suppressed cheat seed produc-
m-2 reduced wheat yield 10, 15, or 20 percent when tion. In the second study (1991), increasing wheat
downy brome emerged within 14 days after wheat seeding rate (from 67 to 101 kg ha1) and decreasing
The Effect of Weed Density 93

row spacing (from 22.5 to 15 cm) decreased cheat filling stage, and Gill et al. (1987) suggested that
density, biomass, and seed harvested with wheat at this diminished the importance of competition for
two of three locations and increased wheat yield. water in wheatripgut brome mixtures. Production
The suppressive effect of wheat was greatest when it of fertile tillers was the wheat-yield trait that was
was seeded in September rather than later in the fall. most sensitive to ripgut brome competition.
McCloskey et al. (1998) studied the interaction of Anderson (1993) showed that jointed goatgrass
a third brome species, poverty brome, with catch- and wheat had similar soil water extraction depths in
weed bedstraw, corn poppy, and wheat. Poverty the U.S. central Great Plains. Jointed goatgrass
brome was the most effective competitor of the four development was similar to the winter wheat culti-
species. Its population increased tenfold under min- var Vona over two seasons with different rainfall.
imum tillage and declined with plowing. Corn The weed at a density of 18 m-2 reduced winter
poppy densities remained low in all trials, and catch- wheat yield 27 or 17 percent when it emerged with
weed bedstraw increased on organically fertilized, wheat or up to 42 days after wheat. Jointed goat-
minimum-tillage plots except when poverty brome grass that emerged in late fall still reduced wheat
was present. High densities of poverty brome yield, but removing jointed goatgrass by early
reduced the population of catchweed bedstraw. In March of the following year prevented wheat yield
the first season of this work (McCloskey et al. loss.
1998), fertilizer had the greatest influence on crop Ogg and Seefeldt (1999) set out to identify the
yield, but subsequently poverty bromes density was competitive traits of seven cultivars of soft white
the most important determinant of crop yield. winter wheat in competition with jointed goatgrass.
Mccloskey et al. (1998) concluded that the interac- The measures were increased wheat yield and
tion of weed species and crop yield was weak. Such reduced jointed goatgrass seed production. In a dry
interactions could be observed but only at high weed year, wheat that grew tall rapidly was able to main-
densities, and the interactions are unlikely to be of tain yield and reduce the weeds seed production. In
great economic importance. a wet year, the number of wheat flower heads per
Pollard (1982) provides a good, but not quantita- plant, the rate of water use, and weight gain were all
tive, box diagram (see fig. 1 of his paper) of the life positively correlated with maintaining wheat yield.
cycle of poverty brome in winter cereals in the UK. Jointed goatgrass seed production was lower in a
Using the diagram, Pollard developed an arithmetic wet year compared to a dry year but Ogg and
model of changes in poverty bromes population. Seefeldt (1999) were not able to identify any culti-
Fleming et al. (1988) included downy brome and var traits correlated with the reduction.
jointed goatgrass in their study of competitive rela- The efficacy of wheat seeding rate in management
tionships with winter wheat. A replacement series of jointed goatgrass in winter wheat was shown in
study was used with all possible combinations of the work by Kappler et al. (2002). Winter wheat grain
three species. In growth chambers with ample fertil- contamination (dockage) was reduced by 6 percent
ity and water and a day/night temperature of for every 10 additional wheat plants m-2 above the
18/10C, the relative total yield of the three species threshold density of 70 wheat plants m-2 at one
was similar, and the authors concluded that they Wyoming location and by only 0.5 percent for each
likely competed for the same resources. Wheat and additional ten wheat plants above a threshold densi-
jointed goatgrass had greater plant growth and a ty of 110 wheat plants m-2 in western Kansas.
higher relative crowding coefficient than downy Increased wheat density reduced jointed goatgrass
brome. Wheat was generally slightly more competi- reproductive tillers in four of six location-year com-
tive than jointed goatgrass at lower soil temperatures binations and biomass in two of four location-year
(18/10C) and higher soil water potentials (-33 kPa). combinations. Kappler et al. (2002) acknowledged
Jointed goatgrass was much more competitive than that the response of jointed goatgrass to wheat den-
downy brome. sity was not consistent over locations or years, but
Gill et al. (1987) reported that ripgut brome was advocated the use of increased wheat density in
an aggressive weed in wheat in Australia, especially jointed goatgrass management programs.
on light soils. They developed an exponential model The first edition of this book reported 17 studies
that adequately described yield loss of wheat due to of wild oat competition in wheat. Wild oat competi-
competition from ripgut brome. The yield loss had tion has been studied for many years in many crops.
been determined before the crop reached the grain- Cereals compete, but not well, with wild oat because
94 Chapter 5

of its normally slower germination. Among the first sity to less than 0.2 m-2 (Philpotts 1975). Without
studies of wild oat competition were those of Pavly- added ammonium phosphate, 70 to 100 wild oats
chenko and Harrington (1935) who demonstrated in yd-2, a density approaching that in Friesens (1960)
careful, now classic, ecological studies that compe- study, were required to reduce yield. Soil fertility
tition began under the soil surface when root sys- (nitrogen status), Bowden and Friesen (1967) sug-
tems mingled and water and nutrients became gested, was a more important determinant of the
limiting. Barley competed more effectively than effect of wild oat on wheat than moderate densities.
wheat because it provided a larger number of semi- These early studies were all reported in the first
nal roots 5 days after emergence and developed edition of this book (Zimdahl 1980) and are included
more crown roots 22 days after emergence than again to make the point that the wild oat problem,
other cereals. Wheats root system was 30 times which has been studied extensively since 1980, is not
larger than that of ball mustard (which depressed new but has occupied the attention of weed scientists
wheat yield up to 40 percent), but wheat was more for many years. Leggett (1983) reported that dockage
severely depressed by wild oat which had a root area due to weed seed in wheat had averaged 2.6 percent
four times greater than wheat. All cereals grown (the range was 2.3 to 4.7 percent) from 1971 to 1981
alone had crown root systems and much larger root at two Canadian terminals. Wild oat had accounted
systems than when grown under the stress of intra- for up to 25 percent of the dockage. Leggett (1983)
or interspecific competition. Cereals grown in 6- assumed the dockage due to wild oat had decreased
inch rows often failed to develop any crown roots. but reported the dockage percentage had remained
Intraspecific competition reduced total root system quite constant. It is safe to assume, as Leggett (1983)
length 81 to 99 times in wheat, rye, and wild oat did, that the wild oat problem has diminished with
when single plants grown in 10-ft squares were time, but the evidence of continued study reported
compared to 18 to 20 plants ft-1 in 6-inch rows here affirms that it has not disappeared.
(Pavlychneko 1937). When wheat was drilled and Gillespie and Nalewaja (1988), working with wild
wild oats or ball mustard were planted between crop oats and wild mustard, showed that wheat yield was
rows, a further six- to tenfold reduction in total root the greatest when both weeds were controlled at or
length was observed. before wheats 2-leaf stage. The weeds were classi-
In early work, Chancellor and Peters (1974) fied as equally competitive. Pollard et al. (1982)
demonstrated that high densities of wild oats are demonstrated the importance of cultural practices in
required to depress yield observably, and that effects work that showed that dicots tended to dominate or
only become visible late in the wheats growth. Wild become dominant when soil was disturbed regularly,
oats affected yield in only 3 of 7 experiments and in whereas perennials and annual grass weeds were
each case at a population greater than 150 plants favored by reduced cultivation. They suggested, and
m-2. No significant yield reduction occurred at 20 to time has verified, that the shift away from plowing
100 wild oat m-2. Friesen (1960) found 135 wild oat will favor annual grass weeds. Pollard and Cussans
yd-2 reduced wheat yield 77.5 percent, compared to (1981) found, consistent with others, that wild oat
14 yd-2. In early studies, Thurston (1962) confirmed was favored by reduced tillage. ODonovan and Shar-
that wild oat can be effectively suppressed by a ma (1983) offered the generalizations related to wild
dense crop of any autumn-sown cereal in the United oats and wheat that increasing seeding rate reduces
Kingdom, but even the densest stand did not com- weed competition, and adding P2O5 at planting, espe-
pletely suppress the weed. The effect was mainly cially with barley, favors the crop over wild oats, but
due to decreased seedling growth. adding nitrogen seems to benefit weed and crop
In Canada, Bowden and Friesen (1967) obtained equally. Morrow and Gealy (1983) provided at least
contrary results because 10 to 40 wild oats yd-2 some justification for the effects of tillage by showing
reduced yields of wheat grown on either summer that wild oat emerged in silt loam from depths up to
fallow land, or on stubble land when ammonium 17.5 cm in a greenhouse study and from 15 cm in the
phosphate was added; also, effects became evident field. There was no emergence from greater depths,
early in the growth. (The rainfall and soil moisture and the greatest emergence was from 5 cm. Thus,
patterns of the relatively wet UK are quite different decreasing tillage will not bring seed near the surface
from dryland Canada.) One Canadian winter fallow where germination is favored.
could reduce wild oat populations by 97 percent. Martin et al. (1987) studied the effects of varying
Two consecutive fallow years reduced wild oat den- wheat density on competition from wild oat and
The Effect of Weed Density 95

winter wild oat. Both species reduced wheat grain and the number of wild oat panicles was reduced.
yield by reducing the number of wheat tillers, but They also attributed this to wheats greater root
the effect could be reduced by increasing wheat competitive ability with delayed wild oat planting.
stand density. When wheat stand density was Shoot competition by wheat was also greater, but the
increased beyond the weed-free optimum was unsat- difference was attributed to root effects.
isfactory for wild oat control. Cousens et al. (1991) used a replacement series
Balyan et al. (1991) investigated the relative com- and found in the UK that wild oats were much slow-
petitive ability and discovered that winter wild oat er to establish than winter wheat or winter barley but
reduced wheat yield 17 to 62 percent depending on had a faster rate of growth after establishment than
the wheat cultivar. Wheat dry matter and grain yield either cereal. In monoculture, wild oats grew more
were not correlated with wild oat dry matter. Wheat slowly than either cereal but were the largest of the
height and dry matter accumulation were reliable three species by the end of the study. The switch
predictors of a cultivars competitive ability but tiller from cereal dominance to wild oat dominance was
number was not. Walker et al. (2002) studied the likely related to wild oats root development, as
effectiveness of competition from three wheat den- Martin and Field (1987) noted, and it occurred late
sities in separate plots in competition with winter in the season, at the flag leaf emergence stage of
wild oat and hood canarygrass. Maximum crop yield cereal growth. Cousens et al. (1991) were not able to
and reduced weed-seed production was achieved for predict, on the basis of their work, that wild oats
hood canarygrass with 80 wheat plants m-2 and a full were always most competitive. In one case, barley
herbicide rate. For winter wild oat, these things were was equally competitive, and in a second location,
achieved with 130 wheat plants m-2 and 75 percent barley was the least competitive of the three.
of the full herbicide rate. The same effects were also The vigor and importance of root competition is
achieved when wheat density was 150 plants m-2 but affirmed in the work of Satorre and Snaydon (1992),
only 50 percent of the full herbicide rate was who demonstrated in box studies that separated root
required. When wheat density was high (150 m-2), and shoot competition, that competition for soil
the full herbicide rate tended to reduce yield partic- resources (especially nitrogen) was more severe
ularly with the herbicides effective against winter than competition for aerial resources (i.e., light).
wild oat. This quite logically also decreased sup- When both root and shoot competition occurred,
pression of weed-seed production. Walker et al. oats and barley were more competitive than wheat,
(2002) strongly advocated that more-competitive but there were significant differences among culti-
wheat cultivars and higher crop densities have the vars of all cereals. Although nitrogen was a major
potential to improve weed control and reduce herbi- factor in competition, nitrogen fertilization did not
cide rates. change the ranking of the cultivars. Cultivars also
Martin and Field (1987) studied competition of differed significantly in shoot competitive ability,
wild oats with wheat in a replacement series and in but these differences were apparently not due to
a separate experiment where the effects on roots and height differences (Satorre and Snaydon 1992).
shoots were separated. In association, the two Henson and Jordan (1982) had previously shown
species competed for the same resources and the that increasing nitrogen fertilization did not elimi-
interaction tended toward mutual exclusivity. Wild nate the depressing effect of wild oats on wheat
oats are more competitive than wheat as Martin and growth or yield.
Field (1987) illustrated by wild oats greater aggres- Cudney et al. (1989) used replacement series and
sivity relative to wheat, their relative yields, and additive series to study wheatwild oat competition.
their shoot dry weights. They attributed wild oats In contrast to Martin and Fields (1987) and
greater competitiveness to their greater root compet- Cousens et al. (1991) work in the UK, Cudney et al.
itive ability. The species were similar in terms of (1989) showed, in California, that in a replacement
shoot competitiveness, but the effects of root and series study, wheat and wild oats were equally com-
shoot competition were additive. Martin and Field petitive, and the yield of wheat grain was linearly
(1988) also showed that wild oats were more com- proportional to the relative density of wild oats. On
petitive than wheat when they were planted simulta- a per plant basis, the shoot dry weight and leaf area
neously and that competitiveness was due to greater index of wild oats were less than those of wheat at
root competition. If wild oats were planted 3 or 6 wheat anthesis. The relative density of wild oats
weeks later than wheat, wheat was most competitive gave a better fit in a regression equation than the
96 Chapter 5

absolute wild oat density. In a separate study, Cud- decrease from 20 to 15C caused a 53-hour delay in
ney et al. (1991) showed that wild oats grew taller the time to reach 50 percent germination of green
and had a greater proportion of their canopy above foxtail seed. Soil water had an even greater effect; at
60 cm at maturity in competition with wheat, thus -650 kPa, green foxtail seed did not germinate at all.
confirming the results of Cousens et al. (1991). In the field, when soil was warm (20 to 25C) and
However, in contrast to Satorre and Snaydon (1992), moist (0 to 400 kPa), green foxtail emerged within a
who found root competition to be most important, few days of wheat. If soil was dry (-400 to -650 kPa)
Cudney et al. (1991) proposed that interference from and temperature was 15 to 20C, green foxtail
wild oats was due to a reduction in wheats leaf area emerged 7 to 14 days after wheat and competition
at early growth stages when wild oats density was was significantly reduced. Wheat normally emerged
low or the plants were not interfering because of the within 6 to 8 days after planting, but green foxtail
distance between small plants. They proposed a took 7 to 21 days to emerge and the differences were
mathematical model to predict the reduction in more pronounced at low temperature and moisture.
wheat growth caused by wild oats. The model is Blackshaw et al. (1981b) clearly illustrated the
based on the light penetration to wheat leaves at effect of time of emergence on competition of green
later growth stages and with higher wild oat density. foxtail in wheat (table 5.4).
For mixed culture the model is: Cool temperatures give early-seeded spring wheat
a competitive advantage over kochia and Russian
thistle (Nord et al. 1999). Fresh weight of wheat was
(LAI %light penetration)
h for mixed culture
greater at 15C than it was for either weed. When
Xt = h=n
growth-chamber temperatures were 23 or 30C,
(LAI %light penetration)
h for mono culture
fresh weights of the weeds were greater than
where, x = ratio of growth rate of wheat in mixed Peterson and Nalewaja (1992) also demonstrated
culture to the growth rate in monoculture, t = time, that yield reductions of hard red spring wheat varied
LAI = leaf area index, and h = each 10-cm with the prevailing environment in field experiments
increment of height (Cudney et al. 1991). conducted over 3 years in North Dakota. Yield
reductions ranged from 0 to 47 percent from 720
Green foxtail competition with wheat has been green foxtail m-2. When yields were regressed based
studied, primarily in Canada, because it was among on weed density alone, the coefficient of determina-
the most prevalent weeds in the 1970s. Yield losses tion was only 0.12. Similar to the work of Black-
ranged from 2 to 25 percent. Green foxtail was not shaw (1981a, b), the coefficient of determination
as competitive as wild oats and wild mustard. Black- improved to 0.62 when multiple regression analysis
shaw et al. (1981a) showed that as few as 100 green included early season temperature, soil texture, and
foxtail m-2 significantly reduced the yield (21 per- foxtail density. Wheat yield reduction decreased as
cent) of a normal height wheat cultivar (Napayo) green foxtail planting and emergence were delayed
and of the semidwarf cultivar Norquay by 44 per- after wheat.
cent. In the same year, 1,600 green foxtail m-2 ODonovan (1994) confirmed the poor relation-
reduced the yield of Napayo 67 percent and ship between weed density or weed dry weight and
Norquay 82 percent. However, in one year (1977) crop yield for competition of green foxtail and pale
1,600 green foxtail m-2 did not reduce yield of the smartweed with wheat, barley, and canola in studies
cultivar Sinton, a cultivar of normal height, but yield over 4 years in Alberta. He suggested this was
was reduced 43 and 54 percent by 800 and 1,200 indicative of the weeds being poor competitors with
green foxtail m-2 in a second year. Thus, the intensi- the crops. If the crops emerged ahead of the weeds
ty of competition was not determined by density and soil moisture was not limiting, yield losses were
alone. There was also no correlation between the minimal.
level of green foxtail competition and the date of The competition of yellow foxtail and barnyard-
seeding. However, Blackshaw et al. (1981b) found grass with barley and spring wheat was studied by
that the soil temperature and soil water at time of Vezina (1992) over 3 years in Quebec. Yellow fox-
planting strongly influenced the early growth of tail infestations of 850 and 240 plants m-2 reduced
green foxtail and were the most important determi- wheat yield in 2 years only when the crop was plant-
nants of the vigor of competition. A soil temperature ed late. Barnyardgrass at 830 to 920 plants m-2
The Effect of Weed Density 97

Table 5.4. The Effect of Time of Emergence of Green Foxtail Relative to That of Two Wheat Cul-
tivars on the Competitive Effect of Green Foxtail on Wheat
Time to 50% emergence Yield reductiona Green foxtail seed
(days) (%) yield (kg ha-1)

Planting date Wheat Green foxtail Sinton Norquay Sinton Norquay

First year
May 24 6 8 12b 22b 161 273
June 2 8 16 12b 15b 59 220
June 17 7 7 6 18 61 73
Second year
May 28 7 13 23b 38b 389 610
June 5 7 1521 7 15b 103 152
June 11 6 16 9 42b 269 472
Wheat yield compared to weed-free plots.
Significant yield reduction, p = 0.05 using Tukeys HSD.

reduced wheat yield in only one year when the crop Canarygrass, Littleseed and Short-spiked
was planted late. When either of the cereals was (Phalaris minor and P. brachystachys)
planted late, the weeds emerged at about the same
time as the crop. But if the crop was planted early Two studies conducted in Greece (Afentouli and
(late April to the first few days of May), the weeds Eleftherohorinos 1996, 1999) showed that the com-
did not affect crop yield. petitive ability of the two species of canarygrass was
The interference of 19 other weeds, most repre- similar but littleseed canarygrass had a faster growth
sented by only a single report, has been studied in rate and more panicles. Wheat yield was not affect-
wheat. Because there is no apparent logic to the ed by a weed density of 76 m-2 but was reduced 36
choice of weed studied by the several researchers to 39 percent by 304 weeds m-2. Neither species
involved, the weeds will be listed alphabetically. affected wheat yield when the weather was cold and
dry during early growth stages. When the weeds
Canada Thistle (Cirsium arvense) were present at harvest, wheat yield was reduced 23
In eight of nine trials over 5 years, wheat yield to 28 percent.
decreased linearly as Canada thistle shoot density Common Lambsquarters (Chenopodium album)
increased (Donald and Khan 1992). There was no
difference in the effect of Canada thistle between A replacement series, conducted in the greenhouse,
no-till and chisel-plowed plots. The effect of Cana- noted that competitive interference for phosphorus
da thistle on wheat yield was greater in years with and, to some extent, for nitrogen was the major limit
more rainfall. In further work (Donald and Khan to wheat growth in the presence of common lambs-
1996), Canada thistle decreased wheat density quarters. Wheat noncompetitively restricted potassi-
(stand) in 3 of 4 years, which was the primary cause um uptake by the weed. Increasing weed density
of reduced wheat yield. The reduction of the number significantly reduced wheat grain size and yield
of wheat spikes per plant and seeds per spike varied (Bhaskar and Vyas 1988).
between years. In contrast, Mamolos and Kalburtji
Common Milkweed (Asclepias syriaca)
(2001) found over 4 years with Canada thistle den-
sities of 0, 4, 16, or 64 plants m-2 that the main fac- In an additive study, wheat yield was reduced 47
tor in wheat yield reduction was nitrogen percent at the highest density of 12 common milk-
concentration. Second, Canada thistle biomass and weed m-2. A simple linear model was as precise as
last its density contributed to wheat yield loss. more-complex models for yield loss prediction
98 Chapter 5

because common milkweed competed late in on summer fallow. Wall (1997) concluded that dog
wheats development (Yenish et al. 1997). mustard was not a vigorous competitor with wheat
or flax.
Common Sunflower (Helianthus annuus)
Field Poppy and Field Violet (Papaver rhoeas
Sunflowers are commonly grown in rotation with
and Viola arvensis)
wheat in the U.S. upper Midwest, consequently vol-
unteer sunflowers frequently are a weed in wheat. Competition between field violet, field poppy, and
When averaged over all planting dates and locations, wheat was studied in two experiments in two succes-
season-long common sunflower competition from sive years in the UK (Wilson et al. 1995). The effects
densities of 3, 9, or 23 plants m-2 reduced wheat of varying crop and weed density were modeled in
yield 11, 19, and 33 percent, respectively (Gillespie terms of weed biomass over time, weed-seed produc-
and Miller 1984). Characteristic of most weeds in tion, and crop yield. Weed biomass declined and a
wheat, common sunflowers were more competitive maximum was reached earlier with increasing crop
when wheat was planted in late than in early May. density. Intraspecific competition was always higher
Wheat yield was reduced 22 percent when 24 com- in the absence of the crop, and it increased with time
mon sunflowers m-2 competed until wheat was in the and weed density. If wheat density (population) was
flag leaf stage. Gillespie and Miller (1984) conclud- halved, the June biomass of field violet increased 74
ed that common sunflower densities of 9 plants m-2 percent and field violet increased 63 percent. Crop
or higher should be removed before the flag leaf yield losses due to increasing weed density were con-
stage to prevent wheat yield loss. sistently greater with low wheat density. Field poppy
was a more vigorous competitor than field violet in
Corn Cockle (Agrostemma githago)
both years. When summer drought restricted late weed
A natural stand of corn cockle had only a negligible growth, wheat yield losses were lower. As crop densi-
effect on barley but reduced winter wheat yields 10 ty decreased, weed-seed production increased to a
percent when crop density was a quarter of normal maximum in weed monoculture.
(Doll et al. 1995). When crop density was normal,
Ivyleaf Speedwell (Veronica hederifolia)
the weed did not affect wheat yield. Doll et al.
(1995) concluded that crop density was more impor- Angonin et al. (1996) demonstrated the importance
tant as a weed management technique than it was for of choosing the right nitrogen fertility level for the
crop yield. Both crops were effective competitors weed to be controlled and for the desired level of
with corn cockle. Each species used growth wheat production. They used three different nitro-
resources better in mixture than when grown alone gen fertility treatments, that is, 60 kg ha-1 at tillering
because the relative yield totals in mixture were plus 80 kg ha-1 at the time of first stem elongation, a
always greater than unity. total of 140 kg ha-1 applied at three times, 60 kg
With 3 years of data, Rydrych (1981) concluded ha-1 applied at tillering, 60 kg ha-1 applied at the first
that if corn cockle was removed monthly from emer- node of stem elongation, and no nitrogen. The com-
gence to harvest, wheats yield decrease was not petitive effects of 17 to 192 weeds m-2 were greatest
significant if the weed was removed before February in the year when ivyleaf speedwell had the best early
in winter wheat that had been seeded the previous growth. Yield losses, described with a nonlinear
October. Corn cockle densities of 170 to 340 plants model, changed each year because of differences in
m-2 lowered wheat yield an average of 18 percent tiller mortality and the variable effects of nitrogen
when competition was eliminated in March. Season- deficiency at stem elongation and flowering. In gen-
long competition, however, lowered yield 60 per- eral, late nitrogen application increased wheat grain
cent, whereas competition only in the fall had no weight and decreased the weeds effects on the
affect on yield. wheat.
Dog Mustard (Erucastrum gallicum) Lanceleaf Sage (Salvia reflexa)
Greenhouse studies suggested that dog mustard was When lanceleaf sage was grown with nitrogen and
less competitive than wheat but had competitiveness phosphorus, it was more competitive than Phalaris
similar to flax. Competition from both crops aquatica. Stress from defoliation or drought
reduced the leaf area, shoot dry weight, height, and adversely affected P. aquaticas competitive ability.
seed production of the weed compared to its growth Lanceleaf sage was more competitive with a sum-
The Effect of Weed Density 99

mer crop of sorghum than with winter wheat (Weer- flax. In 6 of 7 trials, wheat yield loss due to round-
akoon and Lovett 1986). leaved mallow was insignificant. In one trial with
round-leaved mallow density of 237 plants m-2, the
Quackgrass (Elymus repens)
yield loss was only 15 percent. The weed was
Studies in Denmark (Melander 1994) showed that severely suppressed by wheat. Eight weeks after
wheat was less competitive with quackgrass than emergence, the lamina area of round-leaved mallow
rye, but, in contrast to many studies but similar to was less than 3 percent and it produced less than 1
other studies in Denmark, more competitive than percent of the seed of plants growing alone. Wheat
barley. When quackgrass density was 100 primary was so competitive with this mainly recumbent
shoots m-2, the yield loss in rye was about 8 percent weed because wheat reduced photosynthetically
and that in wheat was slightly higher. Yield losses active radiation (PAR) reaching the weed by 80 to
were caused mainly by early competition that 90 percent beginning 4 weeks after crop emergence
caused large reductions in ear number per unit area and lasting for 6 more weeks. In contrast, Makows-
and kernel number per ear, but seed weight reduc- ki (1995) found that round-leaved mallow was com-
tions were not greatly affected. petitive with wheat in two of three experiments.
They used a two-variable model that incorporated
Redstem Filaree (Erodium cicutarium)
early-season crop density loss and round-leaved
Among four crops (oilseed rape, pea, and dry bean), mallow biomass to account for wheat yield loss.
wheat was the most competitive with redstem filaree Losses of up to 60 percent were recorded for wheat
(Blackshaw and Harker 1998). Maximum yield in years and locations where the weed emerged
reduction of 36 percent occurred with redstem fila- before the crop and affected crop emergence. In
ree densities of 100 to 200 plants m-2. Yield progres- wheat, round-leaved mallow density of 200 m-2
sively decreased as the duration of redstem filaree decreased wheat biomass by 100 to 500 g m-2.
interference increased. Three weeks of interference
Russian Thistle (Salsola iberica)
after crop emergence was sufficient to reduce wheat
yield (and the yield of the other three crops). The Young (1988) conducted 2- to 3-year field studies of
mean yield reduction for each week of interference the interference of Russian thistle with winter wheat.
was 1.6 percent for wheat. Based on their results, The wheat yield loss was 0.5 to 0.6 percent for each
Blackshaw and Harker (1998) suggested control of percentage of the total plant biomass contributed by
redstem filaree should be considered and early con- Russian thistle. This was far below the predicted yield
trol is required. A second study by Blackshaw et al. loss of about 10 percent. In one year, rainfall was 46
(2000) determined that redstem filaree was most percent below normal and the highest Russian thistle
competitive with wheat (as nearly all weeds are) density (200 m-2 seeded) produced more than 70 per-
when it emerged before or with wheat and when cent of the total plant biomass and reduced wheat
rainfall in May and June was plentiful. Redstem fila- yield more than 50 percent. The yield was not affect-
ree growth was significantly inhibited by drought. ed until after 6 weeks of interference. However, when
Increasing the wheat seeding rate from 50 to 300 kg rainfall was 65 percent above normal, the same densi-
ha-1 reduced redstem filaree seed production and ty of Russian thistle produced only 20 percent of the
biomass by 53 to 95 percent over 3 years, but wheat total plant biomass and reduced yield only 11 percent.
yield did not increase significantly above a seeding The differences in the effect of Russian thistle
rate of 50 kg ha-1. But, when redstem filaree was between years was related to the rainfall and its pat-
present, increasing wheat seeding rate from 50 to tern, shading by the weed, the time of crop planting,
300 kg ha-1 increased wheat yield from 56 to 498 and the relative time of emergence of the weed and
percent. The increased seeding rate also decreased wheat. In another study, Young (1986) compared the
redstem filaree presence in the soil seedbank by 79 competitive effects of Russian thistle in winter and
percent over 4 years. Thus, similar to several other spring wheat. During the crop growing season, winter
studies reported here, crop seeding rate is an impor- wheat suppressed the weed more than spring wheat.
tant component of integrated weed management. After harvest, the dry weight of Russian thistle that
grew in wheat stubble suppressed up to 75 percent in
Round-leaved Mallow (Malva pusilla)
winter wheat compared to spring wheat stubble.
Friesen et al. (1992) studied the relatively minor Young (1986) concluded that because winter wheat
interference of round-leaved mallow in wheat and reduced seedling establishment, suppressed plant
100 Chapter 5

growth, and reduced the weeds seed production more years, wild mustard seed production was reduced by
than spring wheat, winter wheat should be planted in competition and moisture stress, and seed produced
areas where Russian thistle is known to be a problem. in dry soil was smaller and had negligible dormancy.
One must assume that the wheat played an important Wright et al. (1999) concluded that wild mustards
role, but one must also assume that the season was competitiveness and its potential to produce persis-
responsible for many of the observed effects on the tent seed may be reduced in dry years.
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Vezina, L. 1992. Influence de la date de semis sur la gon State University.
comptition de peuplements de Setaria pumila et
dEchinochloa crus-galli avec lorge et le bl du OTHER SMALL GRAIN CROPS
printemps. Weed Res. 32:5766.
Walker, S. R., R. W. Medd, G. R. Robinson, and B. R. Only five studies of weed interference in other small
Cullis. 2002. Improved management of Avena grain crops were found: two in rye, two in oats, and
ludoviciana and Phalaris paradoxa with more one in a Setaria spp.
densely sown wheat and less herbicide. Weed Res. Downy brome reduced rye yield most when it
42:257270. emerged within 3 weeks of rye emergence but
104 Chapter 5

downy brome densities of more than 100 m-2 were Manthey, F. A., G. A. Hareland, R. K. Zollinger, and
required to reduce rye yield 20 to 30 percent (Black- D. J. Huseby. 1996. Kochia (Kochia scoparia)
shaw 1993). The greatest reduction in rye biomass interference with oat (Avena sativa). Weed Technol.
(28 percent) and seed production (33 percent) 10:522525.
occurred when 400 downy brome m-2 emerged with Melander, B. 1994. Modelling the effects of Elymus
rye. If the same density emerged 6 weeks after rye repens (L.) Gould competition on yield of cereal,
emergence or in early spring (rye is a winter crop in peas, and oilseed rape. Weed Res. 34:99108.
Alberta), biomass and seed yield were reduced less STUDIES OF DIVERSE CROPS
than 10 percent. Winter rye effectively shaded
downy brome thereby reducing its competitive It is well known that forage grasses are susceptible
effect. Among the cereal grains, rye is a vigorous to competition from several other grasses and
competitor. It is more competitive than wheat or bar- broadleaved species. Large crabgrass was more
ley (Melander 1994). competitive than southern sandbur in forage
Kochia interference (30 plants m-2, the highest bermudagrass (Walker et al. 1998). In late season,
density studied) reduced oat yield in 2 of 5 years in without competition, bermudagrass covered 96 per-
North Dakota (Manthey et al. 1996). Kochia inter- cent of the ground. If southern sandbur was present,
ference did not affect oat height, grain test weight, bermudagrass cover was reduced to 81 percent and
groat percentage, or protein content. large crabgrass reduced bermudagrass cover to only
As reported above (see Wheat), no-tillage reduced 72 percent, a significant decrease in forage yield.
wheat population but it did not affect the population Broadleaf dock is perceived as a major problem in
of oat or barley (Lge and Bai 1999). The authors intensively managed permanent grassland in
concluded that no-tillage, compared to tilled man- Switzerland. Niggli et al. (1993) planted young
agement systems, appears to be a good weed man- broadleaf dock into established pure stands of Ital-
agement technique for oat, despite the interference ian ryegrass, perennial ryegrass, meadow foxtail, or
offered by different weed species. Kentucky bluegrass, and studied the effects of cut-
In one experiment, Nandi setaria (Setaria ting interval (every 4 or 6 weeks) and nitrogen (120,
anceps) aboveground dry weight and seed yield 240, or 480 kg ha-1) on broadleaf docks dry matter
was reduced similarly by 20, 40, 80, or 160 production and stem growth. A cutting interval of 6
goosegrass plants m-2. The dry matter yield of 20 to weeks was more favorable to broadleaf dock than
164 goosegrass plants m-2 did not differ signifi- the shorter interval. Nitrogen fertilizer favored the
cantly. If S. anceps density varied from 6 to 9.3 weed, which increased from 2 percent of total
plants m-2, there was little effect on yield of either herbage yield at 120 to 18 percent with 480 kg nitro-
species (Hawton and Drennan 1980). In one exper- gen ha-1. Of the four pasture species, only Italian
iment, when goosegrass emerged 2 weeks or more ryegrass (well known as a vigorous competitor) was
after S. anceps, yield was not affected whether the able to substantially hinder broadleaf docks
crop was planted in rows or broadcast. In a second growth. Niggli et al. (1993) concluded that
experiment, aboveground dry matter of the crop broadleaf dock could not be managed in permanent
was reduced 21 percent when goosegrass emerged pastures of any of the four grasses by competition
2 weeks after the crop. If S. anceps was weeded for from the grasses, cutting or cutting frequency, or
13 days after planting, goosegrass did not reduce nitrogen fertilization.
crop yield. One to three rhizomes of white kyllinga reduced the
shoot fresh weight of two bermudagrass stolons to 56
Literature Cited percent of the control, but green kyllinga did not
Blackshaw, R. E. 1993. Downy brome (Bromus tecto- reduce shoot fresh weight significantly (Kawabata et
rum) interference in winter rye (Secale cereale). al. 1994). Increased planting densities of both kyllinga
Weed Sci. 41:557562. species linearly increased the weeds shot fresh weight
Hawton D., and D. S. H. Drennan. 1980. Studies on and decreased bermudagrasss shoot fresh weight.
competition between Setaria anceps and Eleusine White kyllinga had greater leaf fresh weight, leaf area,
indica. Weed Res. 211215. roots, and rhizomes than green kyllinga, which pro-
Lgre, A., and Y. Bai. 1999. Competitive attributes of duced more shoots and inflorescences.
A. sativa, T aestivum, and H. vulgare are conserved The effect of a natural weed stand on 30 weeded
in no-till cropping systems. Weed Sci. 47:712719. and 30 unweeded 2-year old willow tree plots was
The Effect of Weed Density 105

determined. As one would expect, willow growth in and 80 to 100 percent in system C. There was little
the first year was greater in weeded plots (Sage mortality after 3 years.
1999). However, in the second year willow growth The relative competitive ability of squirreltail (a
was not different between plots, and losses over 2 native perennial range grass) and medusahead (an
years reflected only the effects in the first year of the exotic annual range grass) was measured in a series
study. Soil moisture and nutrient content measured of experiments in western Oregon (Clausnitzer et al.
in midsummer were not different between plots in 1999). Over 3 years, the study included a very dry, a
either year. Because height was greater in weedy dry, and a wetter than normal year. Squirreltail is a
plots and stem number and canopy density were desirable species on rangeland, but the greater inter-
lower in the second year in weedy plots, the authors specific competition of medusahead suggested that
concluded that competition from tall weeds in the it will be difficult if not impossible to establish the
first year was primarily for light. desirable forage species in an existing stand of
Similar to the study of willow, the effects of medusahead unless the weed is controlled. Squir-
herbaceous weeds on loblolly pine grown on plan- reltail seedlings established and grew well without
tations in the southern United States have been medusahead competition.
studied. Seedling tree-height response to weed Gorse, a perennial introduced from Europe, is
control was significantly related to the percentage regarded as a weed in most places, but it was intro-
of the soil covered by weeds, 7 weeks after control duced to Oregon as an ornamental. The possibility
was initiated, and to weed biomass at the end of the of management with perennial ryegrass is shown by
growing season. The primary competition was for the work of Ivens and Mlowe (1980) in New
soil moisture that weeds depleted rapidly (Nelson Zealand. Without cutting, shoot growth of gorse
et al. 1981). Rapid pine growth results when weeds exceeded that of perennial ryegrass over 22 weeks in
are controlled early, but Nelson et al. (1981) were monoculture, but gorse was more inhibited by com-
unable to predict which sites might benefit most petition from perennial ryegrass than the grasss
because of the variable effects of rainfall and vary- growth was reduced by competition from gorse. The
ing levels of weed competition. Britt et al. (1990) gorse root system was small compared to that of
showed that loblolly pine with a low level of weed perennial ryegrass. If both species were cut three
interference (percent ground cover = 1 to 7) had times at 2 to 4 cm, it reduced total growth of both,
more aboveground biomass than trees grown with but gorse was affected more. The implications iden-
a high level of weed interference (percent ground tified by Ivens and Mlowe (1980) are that pasture
cover = 62 to 82). Differences between levels of grasses such as perennial ryegrass should be estab-
weed interference were created by using no con- lished as quickly as possible after gorse clearing to
trol, herbicide in the first year, or herbicide in the limit seedling invasion because a gorse monoculture
second year of growth to control a natural weed will yield more than a perennial ryegrass monocul-
stand. Trees in plots with a low level of weed con- ture. Gorse control will also be aided by grazing ani-
trol were 5 to 10 times larger than those in plots mals.
with a high level of weed interference. Trees grown One study of weed interference in cassava, a
with a high level of weed interference had a lower dietary staple of importance in much of Africa,
percent of their biomass in foliage and a higher Brazil, and India, was found. Giant sensitive plant
percentage allocated to stems (Britt et al. 1990). interference for 12 months at densities of 10,000,
Changes in carbon partitioning with lower alloca- 20,000, 30,000, or 40,000 plants ha-1 and a natural
tion to development of leaf area were suggested to population of 630,000 plants ha-1 was compared. All
be the drivers of accelerated growth associated of the weed populations reduced cassava root yield
with lower weed interference. after 12 months of interference. Yield reduction
After 19 years, ponderosa pine trees were 1.6, 1.9, from the natural population was 85 percent (Alabi et
or 5.7 m tall with no control of bearmat (A), phe- al. 2001).
noxy herbicide control (B), or a combination of her- Oil palm seedlings are commonly started in poly
bicide and clipping (C), respectively (Tappeiner and bags. Initial weeding of the polybags and the sur-
Radosevich 1982). There was a 75 percent net wood rounding area delayed for 16 weeks after planting
reduction after 50 years of bearmat competition. did not decrease seedling growth if weeding every 2
Seedling survival was only 6 to 12 percent in man- to 6 weeks followed (Iremiren 1986). When initial
agement system A, 52 to 88 percent in system B, weeding was delayed until 20 weeks after planting
106 Chapter 5

and subsequent weedings were 8 weeks apart, atropurpureum to intercept more light energy than it
seedling growth decreased, particularly seedling could in monoculture.
height. The authors demonstrated a connection Container-grown ornamentals are valuable and
between weed and disease occurrence by showing normally are weeded regularly to promote good
that when initial weeding occurred 4 or more weeks growth and prevent undesirable transfer of weeds
after planting and the subsequent weeding interval when the plants are purchased and replanted. Few
was longer than 2 weeks, weed growth progressive- studies of the necessity of weed management are
ly increased as did the number of oil palm seedlings done because the necessity does not need to be
affected by Rhizoctonia lamellifera Small and Pythi- demonstrated to nursery owners. Walker and
um splendens Braun. Population was not affected. Williams (1988) showed that barnyardgrass, large
Iremiren (1986) recommended that 4 to 6 weedings crabgrass, and giant foxtail interfered with container-
of oil palm seedings in a poly bag nursery in the 12 grown redosier dogwood as soon as 21 days after
months after planting are as efficient as the normal transplanting. By the end of their study (83 days after
practice of monthly weeding. transplanting), 5 weeds per container reduced
Shoot and root competition of tropical weeds redosier dogwood shoot dry weight as much as 72
from planting until harvest reduced the tuber yield percent. Independent of the number of weeds in a
of white yam 76 to 79 percent, over 3 years of study, container, the same 3 weeds reduced shoot dry
compared to weed-free conditions (Unamma and weight of container-grown bush cinquefoil as much
Akobundu 1989). Without physical contact between as 75 percent after 83 days of interference (Walker
crop and weed roots, leachate from the weed foliage and Williams 1989).
reduced tuber yield 38 to 42 percent, which the
authors noted was clear evidence of allelopathy.
In view of the importance and value of tobacco, it A few studies have been done on the interaction or
is surprising that no studies were reported in the first interference between two weeds. Often these are
edition of this book (Zimdahl 1980), and only one designed to determine why one weed is more com-
has been done since. Presumably because of its petitive and to understand the underlying biology of
value, one is not concerned with whether or not interference. These studies are briefly described
weed control is necessary or when it must be done. below.
It is required. Tobacco yield increased significantly Seven days after planting at 1, 2, 4, or 8 cm, barn-
with weed-free periods of 3 or 4 weeks and yardgrass emerged 96, 90, 83, and 27 percent,
decreased if weed interference lasted more than 3 to whereas redroot pigweed emerged 84, 73, 62, and 0
4 weeks after transplanting (Lolas 1986). As percent from the same depths (Siriwardana and
opposed to the effects of weeds on most crops, when Zimdahl 1984). Intraspecific competition of barn-
tobacco yield decreased, there were also effects on yardgrass was greater than interspecific competition
chemical composition of the crop. from redroot pigweed. When planting depth
The density of Crotalaria goreensis, a weedy increased from 1 to 4 cm and soil moisture increased
tropical legume, when varied from 10, 20, 40, to 200 for 30 to 50 percent (low) to 100 percent (high) of
plants m-2, produced successive reductions in the field capacity, the competitive ability of redroot pig-
yield of Macroptilium atropurpureum, a pasture weed decreased.
crop, and successive increases in C. goreensis yield Lolas and Coble (1980) provided reasons for the
(Hawton and Drennan 1980). When M. atropur- competitiveness of johnsongrass. All growth charac-
pureum density was varied between 2.3 and 6 m-2, teristics (height, leaves per plant, number of tillers,
there was no effect on yield of either species. The fresh weight of rhizomes and shoots) were signifi-
weed lowered the aboveground dry matter yield of cantly increased as the length of planted rhizome
M. atropurpureum only when it emerged 2 weeks segments increased from 2.5 to 25 cm. Johnsongrass
earlier. Removal of C. goreensis 12 days after plant- derived from longer rhizome segments that were
ing was sufficient to prevent a reduction in dry mat- maintained by limited tillage, coupled with the
ter yield of M. atropurpureum. It is interesting to plants rapid growth rate, interfered with crops ear-
note that the species were not mutually exclusive or lier than plants grown from shorter rhizome seg-
harmful, and their mutual effects were always relat- ments.
ed to soil and climate conditions. In mixtures, C. Japanese millet competed well with yellow
goreensis provided physical support and enabled M. nutsedge primarily via root interference (Thullen
The Effect of Weed Density 107

and Keeley 1980). Japanese millet reduced yellow competing for the same resources. Poverty brome
nutsedge dry weight and the number of plants with produced significantly more reproductive tillers and
tubers without any effect on millet. seed after nitrogen was applied and production was
The relative competitiveness of atrazine-resistant greater in the absence of competition.
and atrazine-susceptible populations of common In monoculture, shoot dry weight and leaf area of
lambsquarters and lateflowering goosefoot was spurred anoda and velvetleaf were similar. In mixed
investigated by Warwick and Black (1981). The sus- greenhouse culture, spurred anoda exceeded vel-
ceptible (S) biotype of common lambsquarters out- vetleaf in leaf area per plant and shoot weight (Pat-
competed the resistant (R) biotype, but the biotypes terson 1990). One spurred anoda was equal in
of lateflowering goosefoot were equally competi- competitive ability to 2.5 velvetleaf plants.
tive. Both populations of common lambsquarters Redstem filaree and round-leaved mallow were
had greater total and reproductive biomass and ear- grown in monoculture with 2, 4, 8, or 12 plants in
lier flowering than lateflowering goosefoot. The sus- each 20-cm diameter pot and in all possible combi-
ceptible population of common lambsquarters had nations of equal density. The leaf area per plant was
greater total and reproductive biomass than the similar for both species, but round-leaved mallow
resistant population and was more competitive than grew taller and produced more shoot biomass than
the susceptible population of lateflowering goose- redstem filaree in monoculture. The response in
foot. Weaver and Warwick (1982) did a similar mixed culture varied with the proportion of each
study with redroot pigweed and Powell amaranth. species. When mixed, round-leaved mallow gained
For both species, the susceptible population had leaf area and shoot biomass indicating it was the
greater competitive ability with respect to total bio- superior competitor by about a factor of 2 (Black-
mass and seed production. The susceptible popula- shaw and Schaalje 1993).
tion of Powell amaranth was more competitive than When the competitiveness of wild, dog, and ball
the susceptible or resistant population of redroot mustard was compared, dog mustard had the lowest
pigweed. Redroot pigweeds resistant population leaf area. In a replacement series, wild was the most
was more competitive than Powell amaranths but competitive and greater than ball, which was greater
was about equal to the competitiveness of suscepti- than dog (Wall 1995).
ble redroot pigweed (Weaver and Warwick 1982).
Literature Cited
Perennial ryegrass was more competitive than
white clover when grown in the greenhouse in boxes Alabi, B. S., A. O. Ayeni, A. A. Agbola, and B. A.
that permitted separation of root and shoot competi- Majek. 2001. Giant sensitive plant interference in
tion (Martin and Field 1984). Both can be regarded cassava. Weed Sci. 49:171176.
as weeds but both are also primary components of Blackshaw, R. E., and G. B. Schaalje. 1993. Density
pastoral agriculture in New Zealand. The success of and species proportion effects on interference
the association depends on the nitrogen supplied by between redstem filaree (Erodium cicutarium) and
the white clover and, therefore, maintenance of round-leaved mallow (Malva pusilla). Weed Sci.
white clover is required. Perennial ryegrass is more 41:594599.
competitive than white clover with or without nitro- Britt, J. R., B. R. Zutter, R. J. Mitchell, D. H.
Gjeestad, and J. F. Dickson. 1990. Influence of
gen, at any harvest time. When harvest was 8 weeks
herbaceous interference on growth and biomass
after planting, root competition dominated. For later
partitioning in planted loblolly pine (Pinus taeda).
harvests (16 and 20 weeks), shoot competition dom-
Weed Sci. 38:497503.
inated. For later harvests, nitrogen application Clausnitzer, D. W., M. M. Borman, and D. E. John-
increased perennial ryegrass competition. son. Competition between Elymus elymoides and
The competition of three perennial grasses Taeniatherum caput-medusae. Weed Sci.
(roughstalk bluegrass, poverty brome, and Yorkshire 47:720728.
fog) with poverty brome, used to establish grass Hawton D., and D. S. H. Drennan. 1980. Studies on
strips as field margins, was studied by Rew et al. competition between Macroptilium atropurpureum
(1995) in the UK. Each was grown from seed in and Crotlaria goreensis. Weed Res. 20:225230.
additive mixtures with each species, separately. Iremiren, G. O. 1986. Effect of time of and frequency
There was no significant difference between the rel- of weeding on the growth of polybag oil palm
ative total yield of poverty brome in additive mix- (Elaeis guineensis Jacq) seedlings. Weed Res.
ture with the other grasses indicating they were 26:127132.
108 Chapter 5

Ivens, G. W., and F. Mlowe. 1980. A study of compe- Siriwardana, G. D., and R. L. Zimdahl. 1984. Compe-
tition between seedlings of gorse (Ulex europaeus tition between barnyardgrass (Echinochloa crus-
L) and perennial ryegrass (Lolium perenne L.) by galli) and redroot pigweed (Amaranthus
means of a replacement series experiment. Weed retroflexus). Weed Sci. 32:218222.
Res. 20:183191. Tappeiner, J. C., II, and S. R. Radosevich. 1982.
Kawabata, O., R. K. Nishimoto, and C. Tang. 1994. Effect of bearmat (Chamaebatia foliolosa) on soil
Interference of two kyllinga species (Kyllinga moisture and ponderosa pine (Pinus ponderosa)
nemoralis and Kyllinga brevifolia) on bermudagrass growth. Weed Sci. 30:98101.
(Cynodon dactylon) growth. Weed Technol. Thullen, R. J., and P. E. Keeley. 1980. Competition
8:8386. between yellow nutsedge (Cyperus esculentus) and
Lolas, P. C. 1986. Weed community interference in Japanese millet (Echinchloa crus-galli var. frumen-
burley and oriental tobacco (Nicotiana tabacum). tacea). Weed Sci. 28:2426.
Weed Res. 26:18. Unamma, R. P. A., and I. O. Akobundu. 1989. Effects
Lolas, P. C., and H. D. Coble. 1980. Johnsongrass of tropical weeds on yield in white yam (Discorea
(Sorghum halepense) growth characteristics as rotunda Poir). Weed Res. 29:16.
related to rhizome length. Weed Res. 20:205210. Walker, K. L., and D. J. Williams. 1988. Grass inter-
Martin, M. P. L. D., and R. J. Field. 1984. the nature ference in container-grown Baileys redosier dog-
of competition between perennial ryegrass and wood (Cornus x baileyi) Weed Sci. 36:621624.
white clover. Grass and Forage Sci. 39:247253. . 1989. Annual grass interference in container-
Nelson, L. R., R. C. Pederson, L. L. Autry, S. Dudley, grown bush cinquefoil (Potentilla fruticosa). Weed
and J. D. Walsted. 1981. Impacts of herbaceous Sci. 37:7375.
weeds on loblolly pine plantations. Southern J. Walker, R. H., G. Wehtje, and J. S. Richberg, III.
Appl. Forestry. 5:153158. 1998. Interference and control of large crabgrass
Niggli, U., J. Nsberger, and J. Lehmann, 1993. (Digitaria sanguinalis) and southern sandbur
Effects of nitrogen fertilization and cutting frequen- (Cenchrus echinatus) in forage bermudagrass (Cyn-
cy on the competitive ability and the regrowth odon dactylon). Weed Technol. 12:707711.
capacity of Rumex obtusifolius L. in several grass Wall, D. A. 1995. Comparative analysis of three cru-
swards. Weed Res. 33:131138. ciferous weeds: Growth, development, and compet-
Patterson, D. T. 1990. Effects of density and species itiveness. Weed Sci. 42:7580.
proportion on competition between spurred anoda Warwick, S. I., and L. Black. 1981. The relative com-
(Anoda cristata) and velvetleaf (Abutilon petitiveness of atrazine susceptible and resistant
theophrasti). Weed Sci. 38:351357. populations of Chenopodium album and C.
Rew, L. J., R. J. Froud-Williams, and N. D. Boatman. strictum. Can. J. Bot. 59:689693.
1995. The effect of nitrogen, plant density, and Weaver, S. E., and S. I Warwick. 1982. Competitive
competition between Bromus sterilis and three relationships between atrazine resistant and suscep-
perennial grasses: The implications for boundary. tible populations of Amaranthus retroflexus and A.
Weed Res. 35:363368. Powellii from southern Ontario. New Phytol.
Sage, R. B. 1999. Weed competition in willow cop- 92:131139.
pice crops: The cause and extent of yield losses. Zimdahl, R. L. 1980. Weed-crop competition: A
Weed Res. 39:399412. review. Corvallis, OR: Int. Plant Prot. Center, Ore-
gon State University.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

The Effect of Competition Duration

It is a common but erroneous assumption that After each of these times, the crop is then kept weed
removing weed competition early in the crops free for the rest of the growing season. The second,
growing season, often prior to crop emergence, is frequently complementary, study asks, what is the
the best weed management plan. However, substan- effect when the crop is kept weed free after crop
tial evidence verifies that early weed control, while emergence for certain periods of time and then
not per se wrong, is not essential. The assumption weeds are allowed to grow for the rest of the grow-
that the earlier weeds are removed, the better, may ing season? These studies, when combined, can be
be true for pragmatic reasons such as convenience, used to define what is usually called the critical
combination with other operations, or preparation weed-free period.
for irrigation. Conversely, the assumption may prove Minotti and Sweet (1981) said, weed scientists
false if crop growth and final yield are the operative have conducted a substantial number of so-called
criteria. Unquestionably, the longer weeds compete critical period studies. This review and the first
after crop emergence, the greater their potential edition of this book (Zimdahl 1980) affirm their
effect. However, no effect of any magnitude occurs claim. Minotti and Sweet (1981) note that relatively
(exclusive of allelopathic effects) until competition noncompetitive crops such as onion or garlic require
begins when environmental resources (principally a weed-free period of three months or more. More
water, nutrients, and light) cease meeting the needs competitive crops such as corn or soybeans require
of two or more plants in an area (Clements et al. only three to four weed-free weeks.
1929). Therefore, the mere presence of weeds can- Research to determine the critical period is still
not automatically be judged to be damaging, and it done (35 studies reported here that have been done
does not follow that yield will be reduced if the since 1980) for many crops, and some argue that the
weeds are not controlled immediately. Early in the concept is useful (Evans et al. 2003; van Heemst
growing season, when plants are small, competition 1985). It is useful because one wants to know when
may not occur because the small plants are far apart. weed control should be done. Control is clearly not
When the small plants grow so that they are in close required, for competitive reasons, when both crop
proximity, the competition that occurs, early in the and weed are small. The time when weed manage-
season, is primarily for nutrients and water. Later in ment is performed is determined as much by how
the growing season, when plants are larger, compe- the weeds are to be managed as by the knowledge of
tition may be primarily for light because plants the existence of a critical period. Preemergence her-
shade each other. bicides have been more dominant than they are now,
It seems obvious that if one weed is, or several and they are applied before crop or weed emer-
weeds are, present for 1 day in the life of a crop, it gence, as the label requires, without any considera-
will have no measurable effect on final yield. But tion of the existence of critical period. Now, as
what if the weed(s) is present for 2, 10, or 100 days? Knezevic et al. (2002) point out, the widespread use
The question of duration of competition has been of postemergence herbicides, especially those used
addressed in two ways. The first kind of study asks, with herbicide resistant crops (HRCs), may, but
what is the effect when weeds emerge with the crop there is no guarantee, make the critical period more
and are allowed to grow for defined periods of time? useful and more used. In contrast, van Heemst

110 Chapter 6

(1985) argues that it is not the beginning of the crit- mates of the critical period vary for one crop from
ical period that is important but its end. The end of year to year and site to site. They also showed that
the critical period defines how long weed control the critical period of weed control was different for
must be maintained before crop competition will soybeans grown in a no-till system as opposed to
suffice. Van Heemst (1985) defined the beginning conventional tillage.
and end of the critical period for 16 crops and the In one of the first studies to determine the critical
end of the period for 9 other crops. period, Vega et al. (1967) studied the effect of dura-
Jackson et al. (1985) state, as many others have, tion of weed control on rice. Weeds grew for no time
that the time of weed removal is as important as the at all or in intervals of 10 days up to 50 days after
extent of removal. Weed density is also important. rice was planted. In separate field plots, they also
The greater the weed density, the shorter the time allowed weeds to compete for 10, 20, 30, 40, or 50
the crop can tolerate early-season competition and days after planting and then kept the crop weed free
the longer the required weed-free period will be thereafter. The data (table 6.1) show that yield is
(Weaver et al. 1992). Weaver et al. (1992) also state reduced when rice is weeded for only a short time
that the length of time a crop can tolerate early- after planting. When it was weeded for 40 days,
season competition is related more to the availabili- yield reached a maximum and there was no benefit
ty of soil water and possibly nutrients than it is to from weeding an additional 10 days. In the same
limitations of light. Many critics of the development way, if weeds were allowed to grow up to 20 days
and use of critical periods agree that the length of after planting and then removed, there was no effect
the period (or perhaps its existence) has to vary with on yield. Therefore, rice (and many other crops) can
location and year (Van Acker et al. 1993) and, thus, withstand weed competition early in the growing
a determination of the critical period for a crop- season and does not have to be weeded immediate-
weed combination in one place cannot be valid, ly. On the other hand, weeds in rice cannot be pre-
except as a guide, to what the period may be in other sent more than about 30 days or yield will go down.
places. The point is confirmed by data from Harker With the data from this study, Vega et al. (1967)
et al. (2001) who showed that weed-free pea yields defined the critical period for weed control in rice as
varied between locations by two- to threefold and being between 30 and 40 days after crop emergence.
that affected the length of the critical period Similar data are available for several crops, and
observed. Halford et al. (2001) showed that esti- corn illustrates this. However, the data from the

Table 6.1. The Effect of Duration of Weed Control and Weed Competition on Rice Yield
Weed control duration
(days after planting) Yield kg ha-1

0 46
10 269
20 1,544
30 2,478
40 3,010
50 2,756
Weed competition duration
(days after planting)

10 2,944
20 3,067
30 2,752
40 2,040
50 1,098
Unweeded 55
Source: Vega et al. (1967).
The Effect of Competition Duration 111

many studies do not provide a clear definition of the critical period for weeding. One must assume that
critical period when all studies are considered. the data for each study are accurate but because
Based on the studies reviewed for the first edition of environment and cultural methods play such an
this book (Zimdahl 1980), corn had to be kept weed important role, the data cannot be applied across dif-
free for three to five weeks after seeding or nine ferent geographic regions.
weeks after emergence, dependent on location and These kinds of data have been used to derive the
the weeds. However, when the additional studies critical period for weed competition for many crops.
reviewed for this book are included, the picture is Knezevic et al. (2002) present a complete discussion
not as clear (see tables 6.2 and 6.3). If provided with of the fact that the critical period has been defined in
a weed-free period of three to five weeks after emer- several ways and how the many, often conflicting,
gence, corn will compete effectively with weeds definitions detract from its potential usefulness (Gun-
emerging afterward. Conversely, corn can withstand solus and Buhler 1999). Knezevic et al. (2002) also
weed competition for three to six weeks, if it is then provide a very useful discussion and critique of the
weeded and kept weed free for the remainder of the many ways the critical period has been determined
growing season. These data do not support a precise and make good suggestions for proper statistical

Table 6.2. Weed-free Period Required to Prevent Yield Reduction in Corn

Weed-free weeks required after

Seeding Emergence Competing weeds Location Source

9 Mixed annuals Mexico City Alemn and Nieto 1968

5 Mixed annuals Vera Cruz, Mexico Nieto 1970
3 Giant foxtail Illinois, US Knake and Slife 1969
After 7-leaf stage Redroot pigweed Ontario, Canada Knezevic et al. 1994
3 to 14 leaves Natural stand Ontario, Canada Hall et al. 1992
6 leaves Natural Ontario, Canada Halford et al. 2001

Table 6.3. Length of Early Weed Competition Tolerated without Yield Loss in Corn
Weed-free weeks required after

Seeding Emergence Competing weeds Location Source

3 Mixed annuals Vera Cruz, Mexico Nieto 1970

4 Mixed annuals Mexico City Alemn and Nieto 1968
4 Mixed annuals Chapingo, Mexico Nieto et al. 1968
24 Spreading orach England Bunting and Ludwig
+ Persian speedwell 1964
4 Green foxtail Ontario, Canada Sibuga and Bandeen
6 Giant foxtail Illinois, US Knake and Slife 1969
6 Redroot pigweed Oregon, US Williams 1971
23 Mixed annuals New Jersey, US Li 1960
8 Itchgrass Zimbabwe Thomas and Allison
4 Longspine Colorado, US Anderson 1997
9 to 13 leaves Natural stand Ontario, Canada Halford et al. 2001
14 leaves Natural stand Ontario, Canada Hall et al. 1992
112 Chapter 6

analysis. Generally, the critical period is defined as Although knowing the critical period may have
the time between that period after crop seeding or practical weed management value, Mortimer (1984)
emergence when weed competition does not reduce points out that a limitation is that all weeds are con-
crop yield and the time in the crops life after which sidered equally injurious and no distinction is made
weed presence does not reduce yield. between the kinds of competition that can occur. In
A critical period has been found for several crops. spite of the fact that about 100 studies are identified
As figure 6.1 illustrates, it is a time between the here, few, if any, general principles have emerged.
early weed-free period required and the length of Part of the reason for this is found in the many ways
competition tolerated. Its not a fixed period for any the studies have been done (Knezevic et al. 2002).
crop because it varies with location, season, soil, Perhaps another important reason is that such stud-
and the weeds and their density. Even with its lack ies are relatively easy to conduct, and one rarely
of constancy across environments and seasons, it builds on what has been done. Each is done to deter-
can be a useful measure because it gives evidence mine the critical period, and the only guiding
about when to weed. For example, potatoes, if kept hypothesis is that there must be one. There is a lack
weed free for four to six weeks, will survive the rest of development and testing of hypotheses of mecha-
of the season without yield reduction, even if weeds nism.
grow. If potatoes are weeded nine weeks after seed- The literature reviewed for the first edition of this
ing, yield will not be reduced, if they are subse- book relative to critical periods is included in this
quently kept weed free. Therefore, weeding of edition in the hope that further hypotheses will be
potatoes must be done sometime between four to six developed. These citations have been compiled in
and nine weeks after seeding or yield will decrease. tables 6.4 and 6.5. Some were assembled previously
Critical period analyses show that preemergence by Dawson (1970, 1971). Later, Dawson (1986)
weed control is not essential, nor is weed control generalized that the need for weed control is not
immediately after emergence. This affirms that the based on a number threshold but on a period thresh-
method chosen to control weeds often dictates when old that helps to predict when weed control must be
weeding must be done. One of the useful lessons of applied. The periods are the early-season threshold
critical period studies is that weed control does not or the time after crop emergence before which
have to be done in the first few weeks after crop weeds must be controlled to prevent yield losses.
emergence. Although, for other management rea- This threshold is opposed to what Dawson called the
sons, it often is. late-season threshold or the time after which no

Fig. 6.1. The critical period of competition illustrated for onions. Changes in crop dry weight
from planting to harvest (smooth curve). Yield response from delaying the start of continuous
weeding (); yield response from delaying the termination of weed removal (). (From Mortimer
1984 as adapted from Roberts 1976)
The Effect of Competition Duration 113

further control measures are needed. The data for a the absence of a critical period for weed competi-
particular crop on the presence or length of these tion in red beets (Hewson and Roberts 1973a),
thresholds periods are not very consistent when sev- summer lettuce (Roberts et al. 1977), summer cab-
eral reports are reviewed. Variations in the length of bage (Roberts et al. 1976), and broad beans (Hew-
required weed-free or critical period usually relate son et al. 1973). Ismail (1988) claimed that barley
to differences in competing weeds, geographic grown in Qatar had neither a critical period nor a
region, or the year and environment in which the threshold density below which no yield loss
study was done and illustrate the importance these occurred. Tables 6.4 and 6.5 reveal differences in
factors have. An important difference in the data the periods for these crops, but a single weeding at
developed since 1980 is that a few authors have an intermediate point in time sufficed. However,
specified the percent yield loss that they used to the circumstance varied for onions (Hewson and
determine the critical period. Earlier studies did not Roberts 1973b; Roberts 1973), a crop that has slow
mention the percent yield loss used. It was assumed germination, slow early growth, and susceptibility
that the crop yield was reduced significantly and that to weed competition for a major portion of the
was all that mattered. A few subsequent studies growing season. The extrapolations in tables 6.6
(Amador-Ramrez 2002; Baziramakenga and Ler- and 6.7 also can be questioned on the basis of dif-
oux 1994; Eyherabide and Cendoya 2002; Woolley ferences in competition from specific weeds as
et al. 1993) have introduced the additional factor of illustrated by the data on soybean competition with
the percent yield that counts as a loss. Other studies giant foxtail (Knake and Slife 1969), and tall morn-
have added the crops growth stage rather than just ingglory (Oliver and Schreiber 1973). The giant
time in correct recognition of the fact that crops in foxtail data support the existence of a critical peri-
different environments do not reach the same growth od, but those for tall morningglory do not. This
stage in the same time. Smith et al. (1990) studied emphasizes the importance of each specific weed-
the critical period for velvetleaf interference in cot- crop competition environment and year and that
ton and determined (using only one weed density) conclusions cannot be drawn from one example.
that an inverse linear relationship existed between Additional complications arise when fertility or
velvetleaf dry weight and cotton lint yield but a plant spacing are included as experimental vari-
non-linear equation best described percent lint yield ables, as they ought to be. Li (1960) proposed that
loss as a function of critical-period interference the first 24 weeks after crop emergence were the
interval. most important period of weed competition in corn.
Specific comparisons are difficult when data During this time, weeds completed 16 to 18 percent
cover beans to yams, and range from the West Indies of their total growth, but corn grew only 2 to 3 per-
to Argentina and England. There are some conclu- cent. Weedy corn yield decreased as competition
sions that can be drawn concerning the effect of period lengthened at high fertility, but not at low fer-
competition duration. tility (table 6.8). These data were confirmed by
As stated, the critical period for weed control gen- Bowden and Friesens (1967) study of wild oats in
erally defines the time between when weeds present wheat and flax and Bell and Nalewajas (1968) work
from the beginning of the crop cycle must be with the same weed in barley and wheat.
removed, or the point after which weed growth no Oliver et al. (1976) illustrated the predictable rela-
longer affects crop yield (Nieto et al. 1968; Kneze- tionship between competition duration and weed
vic et al. 2002). Not all studies have been designed spacing. Tall morningglory spaced 15, 30, or 61 cm
to define a critical period; hence, it is not possible to in the soybean row needed 6, 8, and 10 weeks of
decide if such a period exists for every crop or to competition before yield was negatively affected.
know its length. It is reasonable to assume that the Several studies, a few cited below, support the role
difference between the length of weed-free period of plant spacing (Alley 1965; Asberry and Harvey
required (table 6.5) and length of weed competition 1969; Berglund and Nalewaja 1969; Buchanan and
tolerated (table 6.4) represents the critical period, Burns 1971a, b; Dawson 1976; Ivy and Baker 1970;
but a clear division is not present for many crops. Knake and Slife 1962; Mohler 2001; Moolani et al.
The crops for which a critical period may exist are 1964; Smith and Tseng 1970; Smith 1968; Vandiver
shown in tables 6.6 and 6.7. and Wiese 1969; Vesecky et al. 1973; Weatherspoon
The conclusion of the data in tables 6.6 and 6.7 and Schweizer 1971; see chapter 8 for additional
is challenged by work from England that indicates citations).
Table 6.4. Length of Early Weed Competition Tolerated Without Yield Loss by Crops
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Barley 46 leaf stage of crop Wild oats UK Peters 1984

Bean 8 Barnyard grass Washington, US Dawson 1964
Bean 35 after 50% emergence Mixed annuals England Glasgow et al. 1976
Bean 3 Mixed annuals Chapingo, Mexico Nieto et al. 1968
Bean 4 Redroot pigweed Oregon, US Williams 1971
Bean, broad 4 after 50% emergence Mixed annuals England Hewson et al. 1973
Bean, green 4 Purple nutsedge Brazil William and Warren
Bean, lima 6 Sicklepod Georgia, US Glaze and Mullinex
Beets, red 4 after 50% emergence Mixed annuals England Hewson and Roberts
Cabbage 34 Mixed annuals England Roberts et al. 1977
Chapter 6

Cabbage 4 Purple nutsedge Brazil William and Warren

Carrot 5 Redroot pigweed Wisconsin, US Shadbolt and Holm
Acnida sp. 1948
Carrot 3 var. Kuroda; 57 var. Purple nutsedge Brazil William and Warren
Nantes 1975
Corn 3 Mixed annuals Veracruz, Mexico Nieto 1970
Corn 4 Mixed annuals Mexico City Aleman and Nieto
Corn 4 Mixed annuals Chapingo, Mexico Nieto et al. 1968
Corn 24 Spreading orach England Bunting and Ludwig
Persian speedwell 1964
Corn 4 Green foxtail Ontario, Canada Sibuga and Bandeen
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Corn 6 Giant foxtail Illinois, US Knake and Slife

Corn 6 Redroot pigweed Oregon, US Williams 1971
Corn 23 Mixed annuals New Jersey, US Li 1960
Corn 8 Itchgrass Rhodesia Thomas and Allison
Corn 4 Longspine sandbur Colorado, US Anderson 1997
Corn with no-till 913 leaves Unknown Ontario, Canada Halford et al. 2001
Corn 14 leaves Unknown Ontario, Canada Hall et al. 1992
Cotton, winter 17 Mixed annuals Sinaloa, Mexico Ramirez and Nieto
Cotton, spring 9 Mixed annuals Sonora, Mexico Martinez and Nieto
Cotton 8 Mixed annuals Alabama, US Buchanan and Burns
Cotton 2 Mixed annuals India Singh et al. 1971
Cotton 2 Smooth pigweed Rhodesia Thomas and
Apple-of-Peru Schwerzel 1968
Cotton 9 Mixed annuals Arizona, US Arle and Hamilton
The Effect of Competition Duration

Cotton 6 Prickly sida Alabama, US Buchanan et al.
Cotton 4 Yellow nutsedge California, US Keeley and Thullen
Cotton 8.8 Hemp sesbania Mississippi, US Bryson 1990
Cotton 6 Johnsongrass Texas, US Bridges and
Chandler 1987
Cotton 9 Barnyardgrass California, US Keeley and Thullen

Table 6.4. Continued 116

Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Cotton 812 Bermudagrass California, US Keeley and Thullen

Cotton 4 Bermudagrass Georgia, US Vencill et al. 1993
Cucumber 5 Purple nutsedge Brazil William and Warren
Flax 2 Wild oat N. Dakota, US Bell and Nalewaja
Flax 2 Wild oat Manitoba, Canada Bowden and Friesen
Garlic 3 Purple nutsedge Brazil William and Warren
Lentil 4.85.8 Unknown Syria Singh et al. 1996
Lettuce 3 after 50% emerg. Mixed annuals England Roberts and Bond
Chapter 6

Oats 1 Mixed annuals New Jersey, US Li 1960
Okra 3 Purple nutsedge Brazil William and Warren
Oil palm 16 Natural stand Nigeria Iremiron 1986
Onion 4 after 50% emerg. Mixed annuals England Roberts 1976
Onion 4 after 50% emerg. Mixed annuals England Hewson and Roberts
Onion 5 Redroot pigweed Wisconsin, US Shadbolt and Holm
Acnida sp. 1956
Onion 12 Redroot pigweed Nebraska, US Wicks et al. 1973
Annual grasses
Peanut 6 Smooth pigweed Oklahoma, US Hill and Santlemann
Large crabgrass 1969
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Peanut 46 Sicklepod +
beggarweed Alabama, Florida, US Hauser et al. 1975
Peanut 4 Sicklepod +
beggarweed Alabama, Florida, US Buchanan et al. 1976
Peanut 10 Wild poinsettia Georgia, US Bridges et al. 1992
Peanut 2 Common cocklebur Florida, US Royal et al 1997
Peanut 2 Bristly starbur Alabama, US Walker et al. 1989
Seed yield decreased 4%
Peanut 6 to 8 Horsenettle Oklahoma, US Hackett et al. 1987
Peppers, chili 0.73.2 with 5% yield Natural stand Spain Amador-Ramrez
loss deemed acceptable 2002
Potato, sweet 3 Unknown West Indies Kasasian and
Seeyave 1969
Potato, sweet, 4 Natural stand Nigeria Unamma et al. 1985
Maize, and
The Effect of Competition Duration

Potato 46 Unknown Haryana, India Thakral et al. 1989

Potato 4 Goosegrass Java, Indonesia Everaarts and
Torpedograss Satsyati 1977
Smallflower galinsoga
Polygonum nepalense
Potato 6 Redroot pigweed Lebanon Saghir and
Common lambsquarters Markoullis 1974

Table 6.4. Continued
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Potato 2368 w/ high infestation = Quackgrass Qubec, Canada Baziramakenga and

135158 g m-2 Leroux 1994
3 w/medium infestation =
8795 g m-2
15 w/low infestation =
3538 g m-2
Rice, paddy 3 Barnyard grass Arkansas, US Smith 1968
79 Barnyard grass Arkansas, US Smith 1974
4 Ducksalad Arkansas, US Smith 1968
Hemp sesbania
Rice, paddy 6 Smallflower umbrella New South Wales, Swain et al. 1975
sedge Australia
Rice, paddy 3 after transplanting Barnyard grass Philippines Lubigan and Vega
Rice, paddy 4 after transplanting Barnyard grass Philippines Noda 1973
Chapter 6

Rice, paddy 11.4 Spreading dayflower Arkansas, US Smith 1984

Rice, paddy 8 Bearded sprangletop Arkansas, US Carey et al. 1994
Rice, upland 8 Mixed annuals Korea Park and Kim 1971
Rice, upland 6 Mixed annuals Philippines Vega et al. 1967
Rice, wild 7 Common waterplantain Minnesota, US Ransom and Oelke
Sorghum 4 Mixed annuals Nebraska, US Burnside and Wicks
Sorghum 4 Pigweed spp. Texas, US Vandiver and Wiese
Sorghum 6 Tall waterhemp Kansas, US Feltner et al. 1969
Soybean 7 Ivyleaf morningglory Delaware, US Wilson and Cole
Soybean 4 Mixed annuals Illinois, US Wax and Slife 1967
Soybean 89 Giant foxtail Illinois, US Knake and Slife
Soybean 48 dependent upon spacing Tall morningglory Arkansas, US Oliver et al. 1976
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Soybean 24 Sicklepod Alabama, US Thurlow and

Buchanan 1972
Soybean 6 Venice mallow Kansas, US Eaton et al. 1973
Soybean 3 Mustards N. Dakota, US Berglund and
Nalewaja 1971
Soybean 24 Sicklepod Alabama, US Buchanan and
Thurlow 1972
Soybean 8 Common sunflower Missouri, US Allen et al. 2000
Soybean 4 Common cocklebur Arkansas, US Rushing and Oliver
Soybean 4 Mixed, natural stand Illinois, US Jackson et al. 1985
Soybean 4.3 Natural stand Ontario, Canada Van Acker et al.
Soybean 4 Johnsongrass Tennessee, US Williams and Hayes
Soybean 6 Common ragweed North Carolina, US Coble et al. 1981
Soybean, with R-1 = early flowering 810 Natural stand Ontario, Canada Halford et al. 2001
no-till Soybean Giant ragweed Missouri, US Baysinger and Sims
The Effect of Competition Duration

Sugarbeet 12 Barnyard grass Washington, US Dawson 1965

Sugarbeet 4 Kochia Colorado, US Weatherspoon and
Schweizer 1969
Sugarcane 4 Johnsongrass Argentina Arevalo et al. 1977a
Sugarcane 8 Mixed annuals Argentina Arevalo et al. 1977b
Sunflower 4 Mixed annuals Georgia, US Johnson 1971
Tomato 3.4 after transplant Common ragweed Ontario, Canada Friesen 1979
Common lambsquarters
Longspine sandbur
Tomato 3 after transplant Purple nutsedge Brazil William and Warren
Table 6.4. Continued 120
Length (weeks) of competition tolerated after

Crop Seeding Emergence Competing weed(s) Location Source

Tomato 4 after transplant Unknown West Indies Kasasian and

Seeyave 1969
Wheat, spring 2 Wild oat Manitoba, Canada Bowden and
Friesen 1967
Wheat, spring 45 Wild oat England Chancellor and
Peters 1974
Wheat, winter ~22
(Oct., Mar.) Downy brome Oregon, US Rydrych 1974
Yams 12 Unknown West Indies Kasasian and
Seeyave 1969
Yam, white 16 Natural stand Nigeria Akobundu 1981
Chapter 6

Table 6.5. Length of Weed-free Period Required to Prevent Crop Yield Reduction
Weed-free period (weeks) required after

Crop Seeding Emergence Competing weeds Location Source

Barley 2-node stage Wild oat Idaho, US Morishita and Thill 1988
Bean 5 Mixed annuals Washington, US Dawson 1964
Bean, broad 11.5 after 50% emergence Mixed annuals England Hewson et al. 1973a,b
Bean, dry 69 Hairy nightshade Alberta, Canada Blackshaw 1991
Bean, dwarf 11.5 Unknown West Indies Kasasian and Seeyave 1969
Bean, snap Unifoliate stage Common cocklebur New Jersey, US Neary and Majek 1990
Beet, red 24 Mixed annuals England Hewson and Roberts 1973
Cabbage 2 Mixed annuals England Roberts et al. 1977
Corn 9 Mixed annuals Mexico City Aleman and Nieto 1968
Weed-free period (weeks) required after

Crop Seeding Emergence Competing weeds Location Source

Corn 5 Mixed annuals Veracruz, Mexico Nieto 1970

Corn 3 Giant foxtail Illinois, US Knake and Slife 1965
Corn after 7-leaf stage Redroot pigweed Ontario, Canada Knezevic et al. 1994
Corn 314 leaves Natural stand Ontario, Canada Hall et al. 1992
Corn, no-till 6 leaves Natural stand Ontario, Canada Halford et al. 2001
Cotton 6 Mixed annuals Alabama, US Buchanan and Burns 1970
Cotton, spring 4 Mixed annuals Sonora, Mexico Martinez and Nieto 1968
Cotton 4 Johnsongrass Texas, US Bridges and Chandler 1987
Cotton 812 Bermudagrass California, US Keeley and Thullen 1991b
Cotton 7 Bermudagrass Georgia, US Vencill et al. 1993
Cotton 9 Barnyardgrass California, US Keeley and Thullen 1991a
Lentil 12.114.1 Natural stand Syria Singh et al. 1996
Lettuce 3 after 50% emergence Mixed annuals England Roberts and Bond 1975
Pea 1 to 2 Wild oat + tartary Alberta, Canada Harker et al. 2001
Peanut 3 Smooth pigweed Oklahoma, US Hill and Santlemann 1969
Large crabgrass
Peanut 10 Sicklepod, Florida Alabama, US Hauser et al. 1975
The Effect of Competition Duration

Peanut 8 Sicklepod, Florida Alabama, US Buchanan et al. 1976
Peanut 12 Common cocklebur Florida, US Royal et al. 1997
Peanut 6 Bristly starbur Alabama, US Walker et al. 1989
Yield decreased 3%
Peanut 2 Horsenettle Oklahoma, US Hackett et al. 1987
Peppers, chili 0.92.1 after Natural stand Spain Amador-Ramrez 2002
with 5 % 6.715.3 after transplanting
yield loss

Table 6.5. Continued
Weed-free period (weeks) required after

Crop Seeding Emergence Competing weeds Location Source

Potato 9 Redroot pigweed Lebanon Saghir and Markoullis 1974

Common lambsquarters
Rice, paddy 6 after transplanting Barnyardgrass Philippines Lubigan and Vega 1971
Rice, paddy 4.3 Echinochloa spp. Haryana, India Thakral 1989
Rice, paddy 4 Echinochloa spp. California, US Gibson et al. 2002
Rice, upland 3 Mixed annuals and India Sibma et al. 1964
Rice, wild 9 Common waterplantain Minnesota US Ransom and Oelke 1982
Sorghum 3 Mixed annuals Nebraska, US Burnside and Wicks 1967
Sorghum 4 Mixed annuals Nebraska, US Burnside and Wicks 1969
Soybean 4 Tall morningglory Delaware, US Wheatley and Cole 1967
Large crabgrass
Soybean 2 Redroot pigweed Delaware, US Wheatley and Cole 1967
Chapter 6

Soybean 3 Giant foxtail Illinois, US Knake and Slife 1965

Soybean 4 Common cocklebur Mississippi, US Barrentine 1974
Soybean 6 Tall morningglory Arkansas, US Loomis 1958
Soybean 36 Venice mallow Kansas, US Eaton et al. 1976
Prickly sida
Soybean 6 Smooth pigweed Nebraska, US Burnside and Juricek 1967
Tall waterhemp
Green foxtail
Soybean 4 Shattercane Kansas, US Zaresky and Russ 1970
Soybean After V-4 Annuals Balcarce, Argentina Eyherabide and Cendoya
35 2002
130 with 2.5% yield loss
V-0 to V-3 or V-4
1535, V-2 to V-4 stage with
<10% yield loss
Weed-free period (weeks) required after

Crop Seeding Emergence Competing weeds Location Source

Soybean 46 Common sunflower Nebraska, US Irons and Burnside 1982

Soybean, no-till 1st or 2d node Natural stand Ontario, Canada Halford et al. 2001
Soybean 2 (dry year) Common ragweed North Carolina, US Coble et al. 1981
4 (adequate water)
Soybean 810 Giant ragweed Missouri, US Baysinger and Sims 1991
Soybean 4 Johnsongrass Tennessee, US Williams and Hayes 1984
Sugarbeet 6 Kochia Colorado, US Weatherspoon and Schweizer
Sugarbeet 10 Barnyardgrass Washington, US Dawson 1965
Common lambsquarters
Sunflower 46 Mixed annuals Georgia, US Johnson 1971
Tomato, 5 after transplanting Common ragweed Ontario, Canada Friesen 1979
transplanted Common lambsquarters
Longspine sandbur
Watermelon 6 Large crabgrass North Carolina, US Monks and Scholtheis 1998
Wheat, winter 2 Downy brome Nebraska, US Wicks 1966
Yam, white 8 Natural stand Nigeria Akobundu 1981
The Effect of Competition Duration

Table 6.6. Crops with a Critical Period for Weed Competition

Crop Weed-free period requireda Length of competition toleratedb Source

Corn 35 weeks 36 weeks

Potato 46 9 Saghir and Markoullis 1974
Rice, paddy 46 49 Vega et al. 1967
Soybean 24 after planting 48 after planting Several references in tables 6.4 and 6.5
See table 6.5 for supporting data.
See table 6.4 for supporting data.
124 Chapter 6

Table 6.7. Crops with an Identified Critical Period

Crop Critical period Source

Barley infested with wild oat 2-node stage to maturity Morishita and Thill 1998
Bean, snap infested with common Emergence to full bloom of Neary and Majek 1990
cocklebur (Authors note the time snap bean
is too long to be a critical period)
Bean, dry infested hairy 39 weeks after emergence Blackshaw 1991
Bean, white 3% yield loss 2nd trifoliate to 1st flower stage Woolley et al. 1993
Cotton infested with hemp = <62 days after planting Bryson 1990
Cotton infested with johnsongrass 46 weeks after emergence Bridges and Chandler 1987
Cotton infested with barnyardgrass 36 weeks after emergence Keeley and Thullen 1991a
Cotton infested with bermudagrass 47 weeks after emergence Vencill et al. 1993
Intercropped sweet potato, maize 28 days after transplanting Unamma et al. 1985
and cocoyam
Lentil 7.79.3 weeks after emergence Singh et al. 1996
Peanut infested with common 212 weeks after peanut Royal et al. 1997
cocklebur emergence
Peanut infested with bristly starbur 26 weeks after emergence for Walker et al. 1989
tolerated loss of 34%
Peanut infested with horsenettle 26 or 8 weeks after emergence Hackett et al. 1987
Peanut infested with broadleaf 6 weeks after planting, i.e., Chamblee et al. 1982
signalgrass during flowering
Rice infested with bearded sprangletop 2156 days after emergence Carey et al. 1994
Rice, wild infested with common 79 weeks after planting Ransom and Oelke 1982
water plantain
Soybean 938 days after emergence = Van Acker et al. 1993
to 2nd node (V-2) to beginning
pod formation (R-3) stage
Soybean infested with giant ragweed 46 weeks after emergence in Baysinger and Sims 1991
one year and 24 weeks in a
second year
Soybean infested with johnsongrass 45 weeks after emergence Williams and Hayes 1984
Tomato, transplants 2436 days after transplanting Friesen 1979
Watermelon infested with large 06 weeks after emergence Monks and Scholtheis 1998
Yam, white 816 weeks after emergence Akobundu 1981

Few data address the influence of weed planting and maturity decreased progressively as seeding
date on competition because in many studies the date was delayed. Dawson (1976) showed that annu-
weeds studied were those that emerged sponta- al weeds emerging in sugarbeets after July 1 (the last
neously and in other studies weeds are planted and cultivation) were suppressed by the crop and did not
emerge with the crop. Vengris (1963) checked affect yield. Late-emerging weeds were competitive
growth and development of redroot pigweed and in a one-third or one-half stand. Planting prickly
yellow foxtail as affected by time of seeding. The sida or Venice mallow with soybeans reduced yield
earliest seedings produced the tallest plants and 33 percent. When weeds were planted 10 days after
highest weed yield. The interval between emergence soybeans, yield fell 20 percent; weeds planted 20
The Effect of Competition Duration 125

Table 6.8. The Influence of Soil Fertility on Weed Competition in Corn

Yield of weedy plots

Duration of weed competition Low fertility High fertility

(weeks) (bu A) (bu A)

2 111 130
3 114 110
5 114 101
Source: Li (1960).

days after the crop did not affect yield (Eaton et al. regardless of how weeds are to be controlled, and for
1973). Giant foxtail seeded in a band over crop comparing the value of methods with varying per-
rows, three (or more) weeks after corn or soybeans, sistence (Roberts 1976). Importance attaches to the
did not reduce yield of either crop (Knake and Slife specific crop(s) and weed(s) competing and to what
1965). When common lambsquarters was planted 7 resource they compete for under the environmental
days earlier than barley, it could not compete effec- conditions of each study. All of the crops surveyed
tively but when it was planted 21 or 31 days before (tables 6.4 and 6.5) can withstand weed competition
barley, barley could not compete effectively for some duration after planting. Yield-reducing
(Elberse and de Kruyf 1979). The time that tartary competition is likely to occur much earlier in the
buckwheat competed with several crops contributed season, if moisture, rather than light, is the primary
the most to yield loss (de St. Remy and OSullivan limiting parameter (Dawson 1970).
1986). A loss of 0.4 percent per day for wheat, bar- In spite of the abundance of data that purport to
ley, and rapeseed and 1.1 percent per day for oats define a critical period for weed management in sev-
and flax was attributed just to the time the weed was eral different crops, the critical period seems periph-
present in the crop. eral to most weed management decisions. It is clear
Another consideration emerges from the data of that the period varies with the place where the work
Welbank and Witts (1962) who showed that earlier was done and the particular season (wet versus dry,
planting and consequent early weed emergence may hot versus cool, etc.). A casual review of the data
not favor crops because cultivation prior to later assembled here also reveals that the period varies
planting could destroy many seedlings as they with each weed-crop combination, the relative time
emerge. of emergence of the crop and weed(s), that is, the
Kasasian and Seeyave (1969) proposed the work- time competition begins. This agrees with Mohlers
ing hypothesis that crops require a weed-free period (2001) conclusion that dependence of both the tol-
of one-fourth to one-third of their growing period. erated period of infestation and the minimum weed-
Their data confirmed this for beans, tomatoes, sweet free period on a wide variety of factors implies that
potatoes, pigeon peas, sugarcane, and yams. The application of the critical period concept to field sit-
study was based on an earlier paper by Nieto et al. uations requires both extensive data and careful
(1968) reporting that beans and corn were most sus- judgement. Thus, while many critical period stud-
ceptible to weed competition during the first 30 days ies have been done, few have elaborated any gener-
of a 130- to 135-day growth period. The data al principles that permit wide application or
reviewed generally support Kasasian and Seeyaves generalization based on the data generated. Each is
(1969) hypothesis, but with the caveat that useful interesting and, in its own place, perhaps useful to
generalizations still must be confirmed with experi- reinforce what is already knownearly weed con-
mental data. That is, specific weed-crop interactions trol is almost always good compared to late weed
must be considered. control. But, critical period studies have not moved
These concepts and data provide a basis for deter- weed science closer to general hypotheses or new
mining the required duration of weed control, conclusions.
126 Chapter 6

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Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

The Elements of Competition

It is accepted that crop yield reductions are general- already been mentioned in chapter 5. This review
ly, although not always (e.g., when allelopathy is reveals that while the elements of competition are
present), in proportion to the amount of light, water, mentioned frequently, there has been little attempt in
or nutrients weeds use at the expense of the crop the weed science literature to explain the mecha-
(Roush and Radosevich 1985). A very general rule nism of the effect.
is for every unit of weeds grown, there will be one deWit (1960) was among the first to point out the
less unit of crop grown. Inconsistent results between futility, the wrongness, of separating the elements of
weed management experiments in one year or competition. His work advocated a change in the
between years are regularly attributed to environ- approach to the study of competition. He developed
mental (i.e., light, water, nutrient, or climatic) varia- mathematical expressions for competition and advo-
tion. In most cases, data are insufficient to define cated consideration of space and what it contained
cause and effect and the generalization is accepted. rather than studies that separated the components of
For example (one among many), Menges and Tamez competition. For example, competition for light
(1981) stated that soil water (which was not mea- affects growth, which, in turn, affects a plants abil-
sured), soil temperature, and irradiance were more ity to compete for nutrients and water. Competition
useful than weed density to explain the differences will be greatest among similar species that demand
in competition between common sunflower and the same things from the environment. deWits view
onions over two years. No careful measurements of was consistent with current thinking in that species
any factor were made, but the generalization about that best use or first capture environmental resources
the effects is similar to many others. will succeed.
It is simple and neat to study the elements of com- Only in recent years has research begun to con-
petition (nutrients, light, and water) separately. sider the spatial distribution or where weeds are in a
Given the tradition of reductionistic science we have field and how that may affect weed-crop competi-
inherited, it is understandable why such studies are tion. Weed scientists have long been concerned with
done. Reductionism has led to significant scientific what weeds (the species) and how many weeds (the
advances in many fields. It is not wrong to separate density) are present in a field. Control has been
the elements of competition experimentally. Howev- directed at the dominant weed or weeds (see Booth
er, it is wrong to assume that plants separate things, and Swanton 2002, chapter 4). Studies of weed biol-
as science can. It is impossible for plants to separate ogy have emphasized seed production, seed dor-
the elements of competition in nature. As mentioned mancy and survival, and seedling growth,
in chapter 1, plants exist in environments where all establishment, and survival. Results of these good
elements of competition are active and attempts to studies have been translated into management sys-
separate them, while interesting, do not reflect the tems usually without considering the patchiness or
real world of inter- and intraplant competition in nonuniformity of weeds in fields. Control included
which plants exist. the unstated and frequently incorrect assumption
There has been no intent to include every research that weed distribution and density were uniform
report that mentions nutrients, light, or water in this over the field. Thus, tillage or herbicides are nearly
chapter. Most reports included in this chapter have always applied uniformly over a field even though

132 Chapter 7

weed scientists and farmers agree the weeds are not Temperature was the primary factor that con-
distributed uniformly. Farmers and others who try to trolled dominance in competition between Nandi
manage weeds have long recognized that weed dis- setaria and goosegrass over a range of climatic
tribution in a field is not uniform, and control prac- regimes in Queensland, Australia (Hawton 1979,
tices may be reduced or eliminated in some places. 1980). If mean temperatures were less than 23C,
Weed distribution is heterogenous not homogenous. Nandi setaria dominated in mixtures. If mean tem-
Biological knowledge to define how the seed- perature was greater than 23C, goosegrass domi-
bank, seed dispersal, plant demography, and habitat nated (Hawton 1979). Neither species dominated in
interact to determine the stability of weed or weed- the first three weeks of growth. Between three and
seed distribution across fields and across time is six weeks and six and nine weeks, goosegrass was
developing (see Cousens and Mortimer 1995, chap- the superior competitor. Changes in competitive
ter 7). There is a poor understanding of how control advantage were related to growth patterns of the two
techniques affect weed and weed-seed distribution species. Goosegrass dominated at warmer tempera-
over time. As this knowledge develops, weed man- tures, but there was strong indication that at temper-
agers will be able to manage weeds on less than a atures less than 22.1C, low light (an example of the
whole-field basis and that will lead to reduced need interaction of factors) allowed goosegrass to gain a
for tillage and herbicides (Mortensen et al. 1998; competitive advantage (Hawton 1980).
Johnson et al. 1995). The dynamics of patches The time to 50 percent emergence of tomato and
defined as how inherent weed biology interacts spa- four weeds grown at five alternating temperatures
tially with landscape characteristics (Cousens and decreased with increasing temperature and
Mortimer 1995) is an important area of weed man- increased slightly with decreasing soil moisture
agement research. Weed scientists want to under- (Weaver et al. 1988). Base temperatures and thermal
stand why weeds are where they are rather than times required for 50 percent emergence varied
know only what species are present and use the spa- among species but were quite insensitive to soil
tial information as another tool to predict and man- moisture above a critical minimum. Weaver et al.
age weed populations. (1988) suggested that knowing the response of
weeds and crops to temperature could be used as a
management technique to plan optimal planting
It is well known that the differential effects of tem- time and to estimate potential crop yield loss.
perature on photosynthesis and growth influence the Comparison of rate of germination, growth, and
competitive ability of C3 and C4 plants (Christie and development of redroot pigweed, smooth pigweed,
Detling 1982; Pearcy et al. 1981). For example, at and Powell amaranth at day-night temperatures of
any temperature, smooth pigweed (a C4 plant) was 28/22C and 22/14C showed that Powell amaranth
less competitive than common cocklebur or soybean had faster germination and earlier height growth and
(both C3 plants) (Flint and Patterson 1983). There- leaf number than either of the other two species
fore, common cocklebur is more likely than smooth (Weaver 1984). When the three species were mixed
pigweed to compete effectively in early seeded soy- in the field, Powell amaranth had greater competi-
beans. Wheat was a better competitor than jointed tive ability in terms of number of plants, above-
goatgrass at day-night temperatures of 18/10C and ground fresh weight, and seed production.
-33 kPa whereas jointed goatgrass was superior at Patterson et al. (1988) compared the effects of
27/10C and -300 kPa under dry, pacific Northwest temperature and CO2 concentration (350 to 700
conditions (Fleming et al. 1988). Wild oat was more ppm) on cotton, velvetleaf, and spurred anoda
competitive than green foxtail at day-night tempera- grown in growth chambers. Carbon dioxide enrich-
tures of 22/16C than at 28/22C (Wall 1993). The ment decreased the weed-crop ratio for total dry
maximum green foxtail leaf area was at 28/22C weight. This may indicate a competitive advantage
whereas that of wild oat was at 16/10C. Green fox- for cotton with elevated carbon dioxide even if tem-
tails dry weight and leaf area increase occurred ear- peratures are less than optimal for cotton. Cotton dry
lier at higher temperatures. Wild oats leaf area ratio matter increased more (38 percent) than that of
and relative growth rate did not differ between tem- either weed with 700 ppm of CO2 with day-night
peratures (Wall 1993). Green foxtails leaf area ratio temperatures of 26 and 17C. Cottons advantage
was higher, but its relative growth rate was lower at was greater (61 percent) when temperature was 32
lower than at higher temperatures. and 23C.
The Elements of Competition 133

Ziska (2001) found that elevated CO2 significant- vary. The criticism is justified but, in spite of its gen-
ly stimulated leaf photosynthetic rate, leaf area, and eral validity, the weed scientists work has not been
aboveground dry weight of common cocklebur (a C3 without value as pointed out by Radosevich and
plant) more than that of sorghum (a C4 plant) in Roush (1990), who contrasted the quite different but
monoculture. Leaf photosynthesis declined for both equally justifiable objectives of weed scientists and
species when they grew together. However, elevat- ecologists. However, as Tilman (1990) charges,
ed CO2 reduced the percentage decline in common Classical, density-based studies of plant competi-
cocklebur and increased it in sorghum by 35 days tion have demonstrated its existence in nature, but
after sowing relative to ambient CO2. Ziska (2001) have not led to a general theory capable of predict-
proposed that vegetative growth, competition, and ing the dynamics and outcome of plant competi-
potential yield of economically important C4 crops tion.
could be reduced by co-occurring C3 weeds as Several studies that illustrate how nutrients have
atmospheric carbon dioxide increases. been studied by weed scientists follow. Young et al.
(1984) suggested that when light and nutrients were
not limiting, an adequate water supply can eliminate
the effects of quackgrass on corn. High levels of
Patterson (1995) focused on environmental stress in phosphorus or potassium did not overcome quack-
weed-crop interactions. His review included 98 cita- grass interference in soybeans although there was
tions concerning water, nutrients, light, and temper- partial relief by irrigation (Sikkema and Decker
ature. The review also commented on the role of 1987).
increasing levels of atmospheric carbon dioxide in Broomrape infestations in tomato and tobacco
future weed-crop interactions. Patterson (1995) dis- were drastically reduced as higher levels (from none
tinguished between environmental conditions (i.e., to 100 g per pot) of ammonium nitrate (an effect
temperature, wind, soil pH, photoperiod) and envi- above 50 g per pot) or ammonium sulfate (an effect
ronmental resources (i.e., water, nutrients, CO2, and above 60 g per pot) were applied (Abu-Irmaileh
O2). Nearly all of the manuscripts reviewed by Pat- 1981).
terson were included in the first edition of this book Henson and Jordan (1982) showed that wild oat
or are cited here and readers are referred to his competition reduced the effectiveness of nitrate to
review for additional details. increase wheats total plant weight, grain yield, and
Tilman (1990) wrote about the mechanisms of whole-plant percent nitrogen when it was applied
competition for nutrients rather than what happens every four days as K or CaNO3 to pots in the green-
when competition occurs, which has been the appro- house. The findings were confirmed by Carlson and
priate and continuing emphasis in weed science. Hill (1986) who showed that wheat grain yield
Tilmans emphasis was on understanding the decreased with fertilization in wild oat infested
underlying mechanism of competition. Tilman plots. In competition, wild oat was better able to use
acknowledged the extensive ecological and weed added nitrogen, thus gaining a competitive advan-
science literature that affirms the existence of com- tage and reducing wheat yield. Wheat was able to
petition and the equally extensive studies on the respond positively to added nitrogen only when wild
effects of environmental variables on competition. oat density was less than 1.6 percent of total plant
He then quoted Harper (1977, p. 369), it is very density. Their data illustrate the importance of con-
doubtful whether such experiments have contributed sideration of the combined roles of weed density
significantly either to understanding the mechanism and response to nitrogen fertilization. Wheat yield
of competition or to generalizing about its effects. decreases as wild oat density increases and as nitro-
Most of the studies discovered for this review gen fertility increases in the presence of weeds
allude to competition for nutrients but do not go (table 7.1).
much beyond the assertion that it occurs. In Giant foxtail growth in the field and greenhouse
Tilmans (1990) view, this is because the studies increased as nitrogen (from nitrate or ammonium)
have focused on the phenomenon of competition for increased from 56, 112, to 225 kg ha-1 (Salas et al.
nutrients rather than on the mechanism. Phenome- 1997). The weeds total dry weight increased with
nological studies affirm that competition occurs but increasing nitrogen, but seed production reached a
cannot be extended to predict what may happen in a maximum at about 150 kg nitrogen ha-1. Giant fox-
different place, with different species, or as nutrients tail did not show any preference for nitrogen form
134 Chapter 7

Table 7.1. Yield of Wheat Grown in Competition with Wild Oat at Three Nitrogen Levels
Wheat yielda,b
Preplant nitrogen (kg ha-1)

Wild oat density 0 67 134 Averagec

(Plants m-2) kg ha-1

0d 4,280 4,840 4,800 4,640
3 3,990 4,670 5,030 4,560
9 3,230 4,120 3,740 3,700
18 3,560 3,570 2,980 3,370
42 3,280 3,300 2,740 3,107
100 2,860 2,260 2,170 2,429
Averagee 3,533 3,793 3,578
Source: Carlson and Hill (1986).
Average wheat density = 285 plants m-2.
LSD (0.05) wheat yield means = 637.
LSD (0.05) wheat yield means across nitrogen levels = 368.
Control plots were treated with 1.1 kg ha-1 difenzoquat.
Differences among nitrogen level means were not significant.

but seed production decreased when the high rates than light competition) was the primary locus of
were applied as ammonium as opposed to nitrate N. competition between the species.
Salas et al. (1997) concluded that use of ammonium Adding nitrogen fertilizer to apple orchards
fertilizer might be useful as a management tech- increased the nitrogen level in apples but did not
nique to reduce seed production of giant foxtail. completely overcome apple growth inhibition by
Over two years, ivyleaf speedwell decreased Kentucky bluegrass, orchard grass, or Korean les-
wheat ear number per unit area for each nitrogen pedeza (Shribbs et al. 1986).
treatment shown below (Angonin et al. 1996): The relative total yields of mixtures of barley and
beans were significantly greater than 1 when the
Year 1None and 60 kg ha-1 at tillering plus 80 kg
plants root systems were mixed and were reduced
ha-1 at the first node of stem elongation.
when nitrogen fertilizer was applied (Martin and
Year 2None, 60 kg ha-1 at tillering, 60 kg ha-1 at
Snaydon 1982). A conclusion of the study was that
the first node of stem elongation, and 140 kg ha-1
the yield advantage of intercropping in this case was
applied over three dates.
related to the different nitrogen sources used by
The authors explained this by an increase in tiller beans and barley.
mortality and a nitrogen deficiency in wheat at the Blackshaw et al. (2003) studied the response of 23
stem elongation and flowering stages. With late weed species plus wheat and canola to 0, 40, 80, 120,
nitrogen application, individual wheat grain weight or 240 mg kg-1 of soil. Shoot and root growth of all
increased and the weeds effect on wheat yield was species increased with nitrogen rate but the magni-
lowest. tude of the response varied widely among the species
When rice was grown in competition with late studied. Fifteen weeds increased shoot biomass and
watergrass with nitrogen at 0, 60, 120, or 180 kg ha-1, eight increased root biomass more than wheat as N
root dry weight was highly correlated with canopy increased. Ten weeds had shoot biomass increases
structure for both species (Gibson et al. 1999). Late similar to canola whereas five increased root biomass
watergrass showed a significantly stronger response more than canola. All species used more than 80 per-
to nitrogen than rice. When the two plants were cent of available nitrogen at low soil N levels. It is
grown with roots separated or mingled, the results accepted that available evidence shows that added
suggested that root competition for nitrogen (rather fertility affects crop-weed interactions, and that
The Elements of Competition 135

weeds often gain a competitive advantage over crops Phosphorus rates of 0.4, and 0.8 g L-1 of soil with
with added fertility (Ditomaso 1995). Ditomaso veri- plant densities of 2, 4, or 8 per 113 cm2 pots, demon-
fied the common assumption that weeds are more strated that smooth pigweed was not responsive to P
competitive than most crops at higher soil fertility levels but luxurious consumption by the weed
levels, and weeds commonly accumulate higher con- reduced the amount of P available to lettuce and,
centrations of the principal fertility elements (N, P, K, thus, its yield (Santos et al. 1998). Purslane, in con-
Ca, and Mg). It is understood that growers routinely trast, increased in competitiveness in response to P
add nitrogen fertilizer to wheat and canola. It is not but lettuce did not. Santos et al. (1998) concluded
clear how data such as those generated by Blackshaw that P competition appears to be the main mecha-
et al. (2003) can be used to change weed management nism of common purslane interference in lettuce,
practices in either crop. Ditomaso (1995) recom- especially when the crop is grown in low P soil.
mended understanding of the basic mechanisms of Competitive interference for phosphorus and, to
the timing of nutrient uptake by crops and weeds as some extent, for nitrogen was noted between wheat
the necessary means to develop fertilizing strategies and common lambsquarters (Bhaskar and Vyas
that favor crops and disfavor weeds. These strategies 1988). Nutrient interference played a major part in
could include: limitation of growth of wheat plants by common
lambsquarters, whereas wheat exhibited greater
Deep band application
noncompetitive interference in restricting potassium
Nitrification inhibitors
uptake by common lambsquarters.
Intentional shifting of the N sources to ammoni-
Several studies have reported on the concentration
um and urea to restrict growth of ammonium or
of nutrient elements in a crop and interfering weeds.
urea-sensitive weeds
The concentration of N, P, K, and Ca were highest in
Timing of fertilizer application to coincide with
jimsonweed, intermediate in common cocklebur and
specific crop developmental stages and to avoid
tall morningglory, and lowest in large crabgrass
specific weed developmental stages
when all competed in tomato (Sanders et al. 1981).
Conscious selection of crop cultivars
Sanders et al. found a few differences in nutrient
Alteration of crop row spacing or seeding rate to
content in tomato and weed leaf tissue, but there was
increase crop uptake of applied nutrients
no clear relationship between concentration of N, P,
It is clear that future weed management systems K, Ca, Mg, and S and weed density. More fruit was
should consider the effect of rate and time of appli- produced per kg of total assimilated N, P, and K in
cation of fertility on the crop-weed complex. The weed-free than in weedy plots. Bhowmik and Reddy
methods of doing so are not as clear or well devel- (1988) found no difference in nutrient content of
oped. tomato and barnyardgrass in one year, but in a sec-
Kleinig and Noble (1968) said that an important ond year, tomato-leaf N and K levels declined and P
aspect of barnyardgrasss competitive ability is its increased as barnyardgrass density increased.
capability of growing and tillering vigorously early Liebl and Worsham (1987) found that Italian rye-
in the season when light is abundant. Barnyardgrass grass responded better to increasing soil levels of NO3
tillers earlier than rice and addition of superphos- and K than wheat. The net uptake of NO3 and K was
phate stimulates tillering. Final tiller number is also twice as high for Italian ryegrass as it was for wheat.
influenced by nitrogen availability. Barnyardgrass Italian ryegrass responded more to changes in nutri-
competition with rice reduced the number of tillers, ents and had greater ion uptake rates compared to
panicles, and spikelets per panicle. These effects wheat. The weed had greater biomass than wheat
were accentuated by high levels of nitrogen and when grown in monoculture, but due to initial seedling
phosphorus. size, wheat seedlings were larger than Italian ryegrass
Gill and Blacklow (1984) emphasized the impor- seedlings for the first 20 days after emergence.
tance of nitrogen and phosphorus in the interaction In an intercropped culture of corn/cowpea, three
between wheat and great brome. They noted a weeks after planting, weeds in weedy cropped plots
reduction in nitrogen and phosphorus concentration had taken up two to four times as much N, P, K, and
in wheat shoots earlier than in great brome and ear- Ca + Mg as was taken up by the corresponding
lier than significant reduction in dry matter and sug- weed-free crops (Ayeni et al. 1984).
gested than great brome competed with wheat for Ampong-Nyarko and DeDatta (1993) showed that
absorption of both nutrients. in field competition between rice and itchgrass or
136 Chapter 7

junglerice, N availability increased canopy interac- ing light-saturated photosynthesis, leaf respiration
tion light absorption and reduced the leaf area of rates, root:shoot ratio, and leaf density. However, as
rice exposed to sunlight. When no nitrogen was irradiance was reduced from a maximum of 850
applied, there was a difference in dry matter yield mol m-2s-1, support tissues (roots, stems, and peti-
between plants grown at 150 or 400 mol m-2s-1. Dry oles) and leaf area ratios did not change for common
matter increased with increasing light intensity, thus lambsquarters and velvetleaf. These measurements
illustrating the connection between these two ele- increased for soybean and decreased for eastern
ments of competition that plants routinely integrate black nightshade and tumble pigweed. Thus, the lat-
and research separates. We will now turn from nutri- ter species demonstrated superior adaptation for
ents to light as the second of the three major ele- efficient light harvesting in reduced light (Stoller
ments of competition. and Meyers 1989b). The results suggest that chang-
ing plant architecture and the influence of canopy
transmitted PPF may be as important as total dry
matter and leaf area when one tries to describe or
An overview of current research (Holt 1995, 143 predict the effects of crop-weed interference. The
citations) of light effects on plants emphasizes the effect of light deprivation for weed management is
advantages of better understanding of plant shown by work with eastern black nightshade
responses to light quality, transient light, and fluctu- (Stoller and Myers 1989a). When grown in full sun-
ating light environments as a means to manipulate light without interference the weed produced 243 g
the light environment of crop canopies to improve of shoots and 5,957 berries in 20 weeks. Plants
weed management. Holt points out the many grown in 94 percent shade produced only 3 g of
advances made in the ability to measure light that shoots and 23 berries in 20 weeks and 1 g and 1
have implications for weed-crop competition studies berry in 11 weeks. Soybeans effectively, although
and for weed management. Readers are referred to not completely, controlled the weed. The weed pro-
Holts (1995) review for details on light quantity, duced nearly 50,000 seeds per plant in full sunlight
light quality, the effects of changing light environ- but less than 20,000 when grown with soybeans
ments, and the role of transient light. (Stoller and Meyers 1989a). The closer eastern
Aldrich and Kremer (1997) list the several char- black nightshade grew to the soybean row, the lower
acteristics that control plant competition for light in was its productivity and competitiveness.
what they define as the horizontal dimension, which Walker et al. (1988) developed a technique to
is controlled by leaf traits, and the vertical dimen- measure the vertical distribution of leaf area and
sion, a function of plant height. Advantageous leaf thus of light interception within monocultures and
traits include leaf area, leaf angle and arrangement, mixtures of rapeseed, wild mustard, and common
canopy effects (i.e., how deep and effectively light lambsquarters. Light measurements were made at
penetrates the canopy), and light effects within the several levels of the plant canopy and when the
plant community (i.e., shading and competition for information was combined with species height (the
light). vertical component), canopy leaf area index could
The latter point is illustrated in the work of be separated and the contribution of each species
McLachlan et al. (1993b). They asserted that a fun- could be estimated from the sunlit leaf area index of
damental component of modeling crop-weed inter- each species.
ference is the effect of understory photosynthetic It is common, but incorrect, to assume that greater
photon flux (PPF) on weed (and crop) growth. In leaf area will automatically be advantageous for any
other words, it is a measure of how effective shading species. Common lambsquarters has been shown to
is. As PPF declined, dry matter accumulation and be a competitive weed in sugarbeets regardless of
relative dry matter distribution in redroot pigweed where the work was done. When its competitive
was greater in the main stem components than in effects were compared to those of common chick-
branch components. The result was that the propor- weed and sugarbeet, common lambsquarters, the
tion of leaf area and dry matter in the upper parts of worst weed, had the lowest leaf area index (Joenje
redroot pigweed increased as PPF decreased with and Kropff 1987). Its competitiveness is, of course,
increasing corn density (McLachlan et al. 1993b). related to its leaf area but its height, and subsequent
Stoller and Meyers (1989b) found that four weeds competition for light, are more important. If com-
and soybeans adjusted to decreased light by decreas- mon lambsquarters emerged up to 21 days after
The Elements of Competition 137

sugarbeet, the crops yield was still reduced. If the Gossett 1981). Light in the soybean row, three and
weed emerged 30 or more days after the crop, it was five weeks after planting averaged 55 and 40 percent
unable to develop a canopy above the crop and sug- of available light, respectively. This finding was
arbeet yield was not reduced (Joenje and Kropff confirmed in work that showed that shading 30 cm
1987). from the soybean row was similar for all row spac-
The importance of the horizontal determinants ings but the shade inflection point was 15 cm from
cited by Aldrich and Kremer (1997) is affirmed by the row and it occurred more rapidly in 30 as
the work of Barnes et al. (1990). Competition for opposed to 61 or 90 cm rows (Murdock et al. 1986).
light between wheat and wild oat is strongly affect- The study by Jones and Walker (1993) showed a
ed by canopy structure as it influences light inter- linear relationship between light intensity and water
ception and carbon dioxide gain in mixed and pure uptake per unit leaf area over two years. Water
stands. It is good when a plant intercepts as much uptake was proportional to light intensity. Once
light as possible, but to be beneficial the light cap- again we see that while the elements of competition
ture must be translated into carbon fixation and a net can be divided by the reductionistic science, plants
carbon gain. Changes in leaf area and leaf inclina- integrate all things, as they must.
tion affected canopy carbon gain differently in mix- The vigor of competition for light by smooth pig-
tures of the two plants and in monocultures. weed compared to johnsongrass in soybeans was
Competition for light was most influenced by differ- demonstrated by Toler et al. (1996) who showed that
ent positioning of the leaf area in upper canopy lay- smooth pigweed intercepted 2.5 times more light
ers (Barnes et al. 1989). Leaf position in the upper than johnsongrass in one year and 1.8 times more in
canopy layers was the prime determinant of the a second year. In multispecies populations, with 4 or
amount of light intercepted. 8 smooth pigweed 4.6 m-1 of row, light interception
The importance of light capture to competitive by johnsongrass was negligible. Competition for
success is illustrated in several reports. Jones et al. light was the primary reason for soybean yield
(1981) found that an okra-leaf cotton cultivar was decrease caused by hemp sesbania (King and Pur-
less competitive because it had reduced leaf area cell 1997).
compared to cultivars with normal size leaves. The Growth of the vigorous perennial silverleaf night-
fact that redroot pigweed is usually an effective shade is also affected by light. Taproots of plants
competitor was affirmed by growth chamber studies grown in full sunlight had 16 percent greater struc-
(McLachlan et al. 1993b) that showed a linear tural carbohydrates g-1 dry weight than taproots of
increase in redroot pigweeds relative leaf area with plants grown in 92 percent shade. The weeds leaf
temperature over normal growth ranges. The studies area increased in shade but the leaves were thinner
also showed that the weeds relative leaf area was as indicated by the reduced leaf weight per unit area
significantly reduced by canopy induced shading. (Boyd and Murray 1982). Plants grown in 92 per-
Light interference in soybeans has been studied cent shade had 35 percent less chlorophyll per unit
more than in any other crop. Interference between leaf area than unshaded plants but plants grown in
common cocklebur and soybeans was primarily due medium shade (47 percent) had more chlorophyll
to shoot interaction and competition for light (Reg- than plants grown in full sunlight.
nier et al. 1989). When the interference of common Sicklepod, a common weed in soybeans, grew
cocklebur was compared with that of velvetleaf and slightly taller in partial shade, but its dry weight was
jimsonweed in soybeans, common cocklebur was a reduced (Nice et al. 2001). When soybean row width
more effective competitor because at the end of the was decreased from 76 to 38 cm and soybean popu-
growing season it had more leaves in the soybean lation per acre was increased, sicklepod population
canopy, its leaf area was more evenly distributed declined 80 percent, primarily due to light competi-
above and below the soybean canopy, whereas the tion.
other weeds were dominantly above the soybeans, Light has been identified as limiting or, at least,
and common cocklebur had more shade tolerance, an important component in the interaction between
especially in its lower branches (Regnier and Stoller several crops and competing weeds. For example,
1989). Soybean is an effective competitor for light. wild oats limited light penetration and growth of
Maximum shading from the soybean leaf canopy dwarf hard red spring wheat when water and nitro-
occurred 11 weeks after planting and declined 3 gen were nonlimiting (Cudney et al. 1991). The
weeks later due to soybean leaf drop (Murphy and opposite crop-weed relationship was identified
138 Chapter 7

when round-leaved mallow invaded spring wheat in competition, mechanistic explanations have been
Alberta. Round-leaved mallow produced less than 1 elusive. A rice:weed model was developed to evalu-
percent of the seed of plants growing alone because ate the effects of weed leaf area density, leaf angle
wheat reduced photosynthetically active radiation (as leaf light extinction coefficients), and height on
(PAR) that penetrated to the decumbent weed by 80 growth and competition of weeds with rice (Caton et
to 90 percent, beginning four weeks after crop emer- al. 2001). Short weeds and weeds with conical leaf
gence (Friesen et al. 1992). Manipulation of light area densities were weakly competitive, indepen-
has potential to improve wheat yield in fields infest- dent of other traits. For other weed types, interfer-
ed with Italian ryegrass (Ghersa et al. 1994). If total ence was positively correlated with height, and the
radiation reaching the soil surface was reduced to tendency to have more planophile (as opposed to
about 10 percent of full sunlight but the red (600 erectophile) leaves.
nm):far red (730 nm) ratio was maintained at the Corn hybrids with enhanced weed tolerance and
normal ratio of 1.0, wheat grain production declined greater velvetleaf suppressive ability were those
40 percent. When radiation was reduced to 3 percent with a higher leaf area index and a greater ability to
of full sun, and the red:far red ratio was 0.2, wheat capture PAR (Lindquist and Mortensen 1998). The
grain production was 35 percent of that in the con- authors suggested that optimizing corns leaf area
trol. Under all conditions, Italian ryegrass interfer- and, thus, PAR reception would be useful in devel-
ence reduced wheat yield. In full sunlight, the weed oping integrated weed management strategies. In
reduced wheat yield up to 75 percent. Therefore, dry beans, competition for PAR was the principal
shading improved wheat yield in mixed stands and factor in competition with common ragweed
reduced interference from Italian ryegrass. Ghersa et (Chikoye et al. 1996). The ability of common sun-
al. (1994) suggested manipulation of the shade envi- flower to intercept PAR was deemed to be an impor-
ronment could be accomplished by intercropping or tant component of interference in soybeans (Geier et
by strip- or relay-cropping patterns. Similar results al. 1996). A common sunflower density of 0.3 m-2
were obtained in other work. Italian ryegrass had a reduced PAR at the top of the soybean canopy by 24
leaf area index 6.6 times greater than wheat 200 and 18 percent over two years.
days after emergence, and PAR was reduced up to Reductions in tomato yield in Ontario, Canada,
68 percent at wheats boot stage (Hashem et al. were attributed to reduction in light due to shading
1998). Winter rye effectively reduced light to downy by weeds and to weed competition for water
brome by 40 to 90 percent, although the weed still (Weaver and Tan 1987). Reduced light during anthe-
reduced rye biomass and yield (Blackshaw 1993). sis and early fruit set did not affect tomato yield if
Junglerice, goosegrass, and itchgrass had higher light intensity during the rapid fruit development
net CO2 exchange rates at 150, 250, and 400 mol stage was not reduced (McGiffen et al. 1992).
m-2s-1 than rice. Gas exchange rates response to light Some studies have been done to demonstrate the
intensity was greatest during early vegetative stages effect of light level on weed growth independent of
and declined with age. Effects were more evident in crop competition. Flower production in field
rice than in the weeds (Ampong-Naryko et al. bindweed and Russian knapweed declined with
1992). Itchgrass had superior growth and carbon decreasing light. The leaf area of field bindweed
dioxide assimilation than rice, while junglerice and decreased as light decreased from 520 to 325 mol
goosegrass were more susceptible to shading. The m-2s-1, but Russian knapweeds leaf area increased as
major effects of redstem on rice occurred only after light declined from 520 to 236 mol m-2s-1. Dry mat-
the weed grew above the rice canopy, and they were ter of shoots, roots, and rhizomes of field bindweed
attributed to shading that decreased shoot and grain plants grown from seed decreased as light decreased,
production and increased tiller mortality (Caton et whereas plants grown from rhizome segments did not
al. 1997). Caton et al. (2001) acknowledged that produce rhizomes as light decreased. The dry matter
the effects of weed shoot morphology on competi- of Russian knapweed grown from seed or rhizome
tiveness for light in rice have not been well segments decreased as light decreased. For both
described quantitatively and are difficult to study plants, the total PAR was more important than whether
empirically. Their point lends support to the claim low or high light levels occurred first during the study
made at the beginning of this chapter that while (DallArmellina and Zimdahl 1988).
there have been many studies that state an effect of When lower leaves of greenhouse-grown com-
competition due to one or more of the elements of mon cocklebur and velvetleaf were shaded to 5 per-
The Elements of Competition 139

cent of full light for 12 days and upper leaves were pattern of light transmission in the meadow steppe
in full light, lower leaf senescence and leaf area canopy largely accounted for the successful estab-
decreased but branch length and the number of sec- lishment of Kentucky bluegrass on sites with small
ond-order leaves increased. Shading of lower leaves disturbance and little light and the restriction (dom-
increased the leaf area of upper leaves 3 days after inance) of downy brome to sites with more light.
shading began in common cocklebur and 6 days Light played an important role in the dominance
later for velvetleaf (Regnier and Harrison 1993). of triazine-resistant and triazine-susceptible smooth
Total dry weight of velvetleaf 12 days after shading pigweed. When sunlight was 10 percent of full sun-
began was unaffected by shading, but that of com- light, there was no difference in the growth rate of
mon cocklebur was reduced 10 percent. Regnier and triazine-resistant and triazine-susceptible plants
Harrison (1993) concluded that common cocklebur (Ahrens and Stoller 1983). However, with either 100
has greater shade tolerance than velvetleaf but both percent or 40 percent of full sunlight, dry matter
species have the ability to compensate for shading of accumulation 11 weeks after planting was about 40
lower leaves by altering upper shoot growth. percent less in triazine-resistant plants, which may
Yellow nutsedge shoots, number of tubers, height, account for their lack of competitive success in the
and shoot and tuber dry weight were less affected by field.
20, 40, 60, or 80 percent shade than purple nutsedge
(Santos et al. 1997). Shoot and tuber dry biomass of
both species responded linearly to shade. In yellow
nutsedge, 80 percent shade reduced dry matter par- All studies acknowledge the central role of water in
titioning to tubers and increased partitioning to weed-crop interactions. An interesting but apparent-
shoots. In contrast, partitioning to tubers decreased ly ignored approach was recommended by Norris
with 80 percent shade without an increase in parti- (1996) who advocated that water use efficiency be
tioning to shoots. Yellow nutsedge has a lower light used to measure the detrimental effects of weeds and
compensation point that purple nutsedge. The as a way to estimate the cost of weeds. Norris (1996)
authors suggest this may explain the greater world- constructed a graph (fig. 7.1) that compared weed
wide distribution of yellow nutsedge. Yellow occurs biomass in kg dry weight ha-1 and water use in mm
more frequently in regions of low light intensity, ha-1. This method will aid modelers who must con-
whereas purple is more common in tropical areas sider water use and managers who must decide if
with high light intensity. and when to control. Because irrigated land pro-
Artificial shading reduced seed and rhizome pro- duces much of what we consume and water is a
duction of yarrow, and seed production was totally finite and increasing cost resource, it should not be
eliminated at 6.4 percent of full sunlight (Kannan- wasted. Knowing the losses due to weeds and the
gara and Field 1983). The weeds seed production cost of water should allow managers to estimate bet-
was stopped and rhizome production was dimin- ter the need for and benefits of weed control.
ished significantly due to crop interference from It is clear that water does not have a role of equal
barley or peas. magnitude in all crop-weed interactions. For exam-
Bookman and Mack (1983) noted that downy ple, Kropff et al. (1992) showed with a simulation
brome dominates sites of large-scale disturbance model that water shortage only influences the com-
while Kentucky bluegrass dominates sites of small- petitive strength of common lambsquarters when the
scale disturbance in the Festuca (fescue)/Symphori- weed grows above sugarbeets. Otherwise the contri-
carpos (snowberry) habitat of eastern Washington bution of water shortage to competitive interactions
(US). In contrast to many studies in the weed sci- were negligible. The number of days between crop
ence literature, Bookman and Mack (1983) exam- and weed emergence and the temperature in the time
ined the role of light in the relationship and between crop and weed emergence were the most
proposed a mechanistic explanation. Their work important factors affecting competition between
showed that light utilization efficiency of bluegrass common lambsquarters and sugarbeet. In contrast,
was greater although its light compensation point root interference dominated the interactions
was lower than for downy brome. The plants between mayweed chamomile and peas, and soil
respective photosynthetic characteristics (maximum water was more important than nitrogen (Ogg et al.
net photosynthesis was higher for downy brome: 1994). Decreasing soil water potential (-33 to -175
14.9 versus 11.5 mg CO2 dm-2h-1) and the seasonal kPa) reduced several aspects of pea growth and
140 Chapter 7

Fig. 7.1. Relationship between water used and weed biomass production. The equation is mm
ha-1 = coeff* x/1000, where coeff is the transpiration-coefficient for either C3 (666) or C4 (300)
plants, and x is the plant biomass in kg dw ha-1. (Reprinted with permission of the Editor, Weed
Technology [Norris 1996])

increased the aggressiveness of mayweed with soybeans increased water uptake per plant and
chamomile toward pea. per unit leaf area (Jones and Walker 1993), but the
Similarly, well-watered and drought-stressed effects were not identical for common cocklebur and
common cocklebur reduced soybean yield 29 versus sicklepod. Canopy interference by soybeans with
12 percent (Mortensen and Coble 1989). Drought- sicklepod increased the soybeans water uptake per
stressed common cocklebur interfered with soybean unit leaf area. Root interference by soybeans
over a shorter distance and the magnitude of the decreased water uptake per plant by common cock-
effect at any distance was reduced. Soybeans yield lebur and root and canopy interference by soybeans
potential was reduced by water stress and drought decreased water uptake by sicklepod (Jones and
caused a reduction in common cocklebur growth. Walker 1993). The leaf area and shoot weight of all
Interference between soybean and common cockle- three species decreased as a result of root interfer-
bur was not stable across all soil-moisture condi- ence from any other species. Common cockleburs
tions, which has implications for modeling efforts. water uptake was twice that of soybeans or sickle-
Midday xylem potentials of soybeans and com- pod
mon cocklebur decreased as the growing season pro- Soybean leaf area and aboveground biomass were
gressed and were lower in common cocklebur greater than those of Florida beggarweed under opti-
during the soybeans vegetative and reproductive mum soil moisture conditions, but they were equal
phases. The differences in xylem potential between to or less than the weed under water stress (Griffin
soybeans and common cocklebur on a given day et al. 1989). Soybeans were more competitive with
were small (Scott and Geddes 1979). Greater diffu- adequate soil moisture but less so with drought
sive resistance was more common in soybeans than stress. As water stress increased, stomatal conduc-
in common cocklebur and, for each species, under tance, photosynthetic rate, and transpiration of vel-
the stress of competition. Canopy interference and vetleaf declined more rapidly than they did in
canopy and root interference of common cocklebur soybeans (Munger et al. 1987). It is clear, perhaps
The Elements of Competition 141

the only clarity, that water stress does not have an only from the upper 75 cm of the soil profile (Cast-
equal effect on crops and weeds. ner et al. 1989).
Among seven weeds that compete with soybeans, Wheat yield decreased more from Canada thistle
the net photosynthetic rate, net assimilation rate, and competition in years with higher rainfall (Donald
water use efficiency on a whole plant or a single leaf and Khan 1992). Jointed goatgrass seed production
basis were greatest in C4 smooth pigweed (Patterson was lower in a wet than in a dry year (Ogg and
and Flint 1983). Smooth pigweed affected cotton Seefeldt 1999). The number of wheat heads per
water relations by reducing plant water stress early plant, wheats rate of water use, and its weight gain
in the season and by shading late in the season (Stu- were positively correlated with maintaining yield in
art et al. 1984). Smooth pigweed has the capacity to a wet versus a dry year, but no firm relationships
extract water from lower in the soil profile, it also were established between the growth of wheat and
had higher diffusive resistance and reduced transpi- jointed goatgrass and moisture supply.
rational losses. The predicted yield loss was similar under normal
Growth reduction associated with water stress water conditions when common cocklebur compet-
was greater in soybeans than in sicklepod (Patterson ed with peanut (Royal et al. 1997). When the water
1986). With adequate water, competition from sick- supply was above normal, peanut yield was 9 to 24
lepod decreased the soybeans leaf area duration, but percent lower under competition.
competition from soybeans decreased the weeds Reductions in tomato yield due to weed interfer-
leaf area duration and net assimilation rate. Thus, ence were attributed to shading by the weeds (as
the effects were similar but the magnitude of the mentioned above) and to competition for water,
effect was greater on soybeans (Patterson 1986). which resulted in stomatal closure (Weaver and Tan
No competition for water or light was detected in 1987).
a study of ivyleaf morningglory and soybeans
(Cordes and Bauman 1984). On the other hand, the COMPETITION FOR OTHER
yield of quackgrass infested soybeans was increased ENVIRONMENTAL FACTORS
by irrigation, although irrigation did not eliminate Clements et al. (1929) limited competition primari-
all effects of quackgrass interference (Young et al. ly to nutrients, water, light, and perhaps space. How-
1983). ever, plants require other factors for growth, but
Patterson and Highsmith (1989) showed that competition for these has not received extensive
water stress reduced cottons height, total dry study. Isolating specific affects is difficult, plus the
weight, and leaf area in competition with velvetleaf primary factors are so dominant in the environment
or spurred anoda when compared to well-watered and relatively easy to isolate for study. The first edi-
controls. Drought did not affect the relative compet- tion of this book (Zimdahl 1980) cited only 11 stud-
itive abilities of the three species or the weeds ies that dealt with factors other than nutrients, light,
effects on cotton. The weeds effects were apparent or water, and four were about temperature, which is
as early as 11 days after the onset of competition. covered above. No additional studies that carefully
Bermudagrass significantly reduced soil-water defined the actual or potential role of other environ-
content 15 cm deep, but soil water was not affected mental factors (e.g., soil microsite difference, soil
30, 45, or 60 cm in the soil profile in competition pH, soil atmosphere, and carbon dioxide) were
with cotton (Vencill et al. 1993). found in the weed science literature.
Volumetric water content up to 180 cm deep in the
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vetleaf (Abutilon theophrasti) with cotton Weed Sci. 46:307312.
(Gossypium hirsutum) during simulated drought Santos, B. M., J. P. Morales-Payan, W. M. Stall, T. A.
and recovery. Weed Sci. 37:658664. Bewick, and D. G. Shilling. 1997. Effects of shad-
The Elements of Competition 145

ing on the growth of nutsedges (Cyperus spp.). Vencill, W. K., L. J. Giraudo, and G. W. Langdale.
Weed Sci. 45:670673. 1993. Response of cotton (Gossypium hirsutum) to
Scott, H. D., and R. D. Geddes. 1979. Plant water coastal bermudagrass (Cynodon dactylon) density
stress of soybean (Glycine max) and common cock- in a no-tillage system. Weed Sci. 40:455459.
lebur (Xanthium pensylvanicum): A comparison Walker, G. K., R. E. Blackshaw, and J. Dekker. 1988.
under field conditions. Weed Sci. 27:285289. Leaf area and competition for light between plant
Shribbs, J. M., W. A. Skroch, and T. J. Monaco. 1986. species using direct sunlight transmission. Weed
Interference between apple (Malus domestica) Technol. 2:159165.
seedlings and four ground cover species under Wall, D. A. 1993. Comparison of green foxtail
greenhouse conditions. Weed Sci. 34:533537. (Setaria viridis) and wild oat (Avena fatua) growth,
Sikkema, P. H., and J. Dekker. 1987. Use of infrared development, and competitiveness under three tem-
thermometry in determining critical stress periods perature regimes. Weed Sci. 41:369378.
induced by quackgrass (Agropyron repens) in soy- Weaver, S. E. 1984. Differential growth and competi-
beans (Glycine max). Weed Sci. 35:784791. tive ability of Amaranthus retroflexus, A. powelli,
Stoller, E. W., and R. A. Meyers. 1989a. Effects of and A. hybridus. Can. J. Plant Sci. 64:715724.
shading and soybean (Glycine max L.) interference Weaver, S. E., and C. S. Tan. 1987. Critical period of
on Solanum ptycanthum (Dun.) (eastern black weed interference in field-seeded tomatoes and its
nightshade) growth and development. Weed Res. relation to water stress and shading. Can. J. Plant
29:307316. Sci. 67:573581.
. 1989b. Response of soybeans (Glycine max) Weaver, S. E., C. S. Tan, and P. Brain. 1988. Effect of
and four broadleaved weeds to reduced irradiance. temperature and soil moisture on time of emer-
Weed Sci. 37:570574. gence of tomatoes and four weed species. Can. J.
Stuart, B. L., S. K. Harrison, J. R. Abernathy, D. R. Plant Sci. 68:877886.
Krieg, and C. W. Wendt. 1984. The response of cot- Young, F. L., D. L. Wyse, and R. J. Jones. 1983.
ton (Gossypium hirsutum) water relations to smooth Effect of irrigation on quackgrass (Agropyron
pigweed (Amaranthus hybridus) competition. Weed repens) interference in soybeans (Glycine max).
Sci. 32:126132. Weed Sci. 31:720727.
Tilman, D. 1990. Mechanisms of plant competition . 1984. Quackgrass (Agropyron repens) inter-
for nutrients: The elements of a predictive theory of ference in corn (Zea mays). Weed Sci. 32:226234.
competition. In Perspectives on plant competition, Zimdahl, R. L. 1980. Weed/crop competition: A
ed. J. B. Grace and D. Tilman, 117141. San review. Corvallis, OR: Int. Plant Prot. Center, Ore-
Diego, CA: Academic Press. gon State University.
Toler, J. E., J. B. Guice, and E. C. Murdock, 1996. Ziska, L. H. 2001. Changes in competitive ability
Interference between johnsongrass (Sorghum between a C4 crop and a C3 weed with elevated car-
halepense), smooth pigweed (Amaranthus bon dioxide. Weed Sci. 49:622627.
hybridus), and soybean (Glycine max). Weed Sci.
Weed-Crop Competition: A Review, Second Edition
Robert L. Zimdahl
Copyright 2004 by Blackwell Publishing Ltd

Weed Management Using
the Principles of Competition

Perhaps, to use the current vernacular, this chapter is Liebman and Dyck (1993b) acknowledged the seri-
where the rubber meets the road. We must ask, has all ous problems with conventional weed management
the work that has been done to establish that weeds strategies and proposed that ecologically based
affect crop yield changed weed management? Has the alternatives ought to be examined and tested. I
abundant information improved weed management? have expressed similar concerns (Zimdahl 1998a, b,
One view (Norris 1992, 1999), based on an extensive 1999, 2002; Zimdahl and Speer 2001). Liebman and
survey (Norris 1997), concluded that in spite of the Dyck (1993b) offered five reasons in support of the
abundant literature on the effects of weed density and need to develop ecologically based weed manage-
duration on competition, improved computer technol- ment systems:
ogy, and decision-aid models, the information on
1. Herbicides have undesirable effects on the
weed-crop competition has had almost no effect on
quality of surface and groundwater.
weed management practice. Norris (1999) strongly
2. Many herbicides are becoming less effective
argued for greater emphasis on weed biology and
due to development of herbicide-resistant
research to understand the mechanisms of competi-
biotypes (Heap 2003).
tion. The evidence in this review is that his plea has
3. Many herbicides are being removed from the
not resulted in a significant change in the research
market due to declining sales, increasing reg-
weed scientists do.
ulatory requirements, or herbicide resistance.
It is not that the dominant control orientation has
4. Herbicides may not be a viable option for
failed to fulfill its goalto control weeds in a crop
farmers in developing countries and are not
and prevent yield loss. It is that the science has not
an option for those who wish to farm organi-
moved beyond the ability to control nearly all weeds
selectively in nearly all crops. That ability is not a
5. Ecologically based weed management can
trivial achievement. It is an achievement that took
be part of agronomically productive, econom-
the combined efforts of university research scien-
ically viable farming systems.
tists, cooperative extension specialists, and chemical
industry scientists. They are rightfully proud of the No one has demonstrated that the reasons offered
progress that has been made from the days of the by Liebman and Dyck (1993b) are bad or that devel-
hoe and horse to the present era of selective, eco- opment of ecologically based weed management
nomical, relatively safe, efficient weed control. systems is a bad idea. Little research has been done
However, as successful as these methods are, they to develop such systems in comparison to the great
have also created problems that have been ignored emphasis on work to preserve the present produc-
or dismissed as externalities1 for too long. These tion system. Little of the work that has been done to
serious problems include harm to nontarget species, describe the consequences of weed-crop compe-
harm to humans, environmental harm, ground and tition has led to development of the required prin-
surface water pollution, soil pollution, and often ciples to develop ecologically based weed
high cost. The undeniable success of modern, chem- management systems. The challenge remains.
ically, energy, and capital intensive agriculture has The remainder of this chapter is divided into eight
also discouraged investigation of other options. parts that reflect the research done since 1980: (1)

Weed Management Using the Principles of Competition 147

plant arrangement in the community, (2) monocul- For four soybean cultivars, there was little effect
ture versus polyculture, (3) tillage, (4) rotation or of changing row widths on soybean yield if the crop
crop sequence, (5) shade, (6) the role of crop geno- was planted on what Parker et al. (1981) called nor-
type, (7) fertility, and (8) the importance of weed mal dates. If planting was later than June, soybean
biology and ecology. yield decreased as row width increased. In other
work, Costa et al. (1980) found that among several
soybean cultivars, early-maturing cultivars had a
greater yield response (+27 percent) to narrow (27
As first reported in chapter 5, Mohler (2001) cm) rows than did cultivars in maturity groups I and
claimed that the density, arrangement, cultivar, and II (+19 percent). Soybean cultivars grown in 27 cm
planting date of the crop that maximizes the rate at rows versus the more conventional 76 cm rows pro-
which the crop occupies space early in the growing duced an average seed yield 21 percent higher over
season usually minimize competitive pressure of all years, populations, and cultivars (Costa et al.
weeds. Mohler reviewed 91 papers that dealt with 1980). Late-maturing cultivars produced the highest
29 crops and only found 6 papers that failed to ver- yields over all row spacings, plant populations, and
ify that increasing crop density resulted in decreas- years. Van Acker et al. (1993) advocated narrow
ing weediness. Mohlers (2001) review also found rows or early-branching soybean cultivars to reduce
that at any given crop density, the slope of the weed weed competition.
biomass curve is greater when density of the weed is Over two years, as row spacing increased, weed
high. He concluded and provided data to verify that resurgence (growth after initial control) increased
suppression of weeds and increase in crop yield (Yelverton and Coble 1991), and the growth was
from an incremental increase in crop density directly correlated with the amount of light pene-
increases with the density of weeds. trating through the soybean canopy to the soil sur-
Most of the work reported in this section deals face. In one experiment (Esbenshade et al. 2001),
with the effects of row spacing on weeds and most row spacing (38 versus 76 cm) had no effect on bur-
(nine reports) was done in soybeans and nearly all cucumber emergence or biomass production.
of those studies and those on other crops show that Sicklepod growth was less in narrow (25 cm) than
weed populations and their effects were reduced as in wide (102 cm) rows (McWhorter and Sciumbato
row width decreased. In view of the early agro- 1988). Reducing soybean rows from 76 to 38 or 19
nomic work on optimum seeding rates (e.g., see cm while increasing soybean population reduced
Martin et al. 1976) and on the effects of row spac- sicklepod population up to 80 percent. Smith and
ing on crop yield, the effects of row width on Jordan (1993) showed that sicklepod growth and
weeds is not surprising, although narrow rows do morphology responded to its distance from the soy-
not always result in reducing the effects of weeds. bean row and its time of emergence relative to the
With crops, narrower rows tend to optimize yield crop. Virtually all sicklepod plants were taller than
until intraspecific interference increases and crop soybeans, but if the weed grew close to the soybean
yield decreases. When crops and weeds interfere, row, its height, number of main stem nodes, number
one may strive to optimize crop yield by employ- of primary branches, and shoot dry weight all were
ing narrow rows to minimize the weeds detrimen- lower. Sicklepods shoot dry weight could be
tal effects. The effects of higher crop populations reduced up to 60 percent if it grew close to the soy-
(narrower row widths) usually can be expected to bean row (Smith and Jordan 1993). In most cases,
reduce weed competition. But that is not always proximity to the soybean row also reduced sicklepod
true. In cabbage and cucumbers, narrowing row seed production (Nice et al. 2001). In competition
width resulted in smaller crop plants and a shorter with sicklepod, it was only under optimal growing
time during which the crops could remain weed conditions (in one year) with conventional cultivars
infested without diminishing yield (Weaver 1984). and sequential herbicide application that 19 cm rows
A point confirmed by Mohlers (2001) observation produced more yield (21 percent) than 38 cm rows
is that the effects of intraspecific competition fre- with medium soybean populations (455,375 plants
quently depress harvest index when the crop is ha-1) in Mississippi (Buehring et al. 2002). Further
grown at high densities. This effect will, of illustrating the interaction of row width and plant
course, vary with the biology of the crop and the population, Buehring et al. (2002) showed that with
interfering weed(s) (Mohler 2001). a low soybean population (241,000 plants ha-1)
148 Chapter 8

19 cm rows yielded 64 percent more than 76 cm increasing wheat planting rate from 67 kg to 134 kg
rows. However, in a second year, with similar sick- ha-1 reduced rye seed production 21 and 25 percent in
lepod control, there was no difference in yield two experiments. At one site, doubling wheat seeding
between 19 cm and 38 cm rows. However, in 19 cm rate in 10 cm and 20 cm rows increased the yield of
rows, soybean yield was 15 percent higher and in 38 rye infested wheat 23 to 27 percent, but there was no
cm rows it was 24 percent higher than yield in 76 cm benefit in 30 cm rows (Roberts et al. 2001). Increasing
rows (Buehring et al. 2002). Although row spacing wheat density up to 800 plants m-2 reduced Italian rye-
did not affect soybean height or seed size, the num- grass seed yield 87 percent but increased its harvest
ber of pods was higher in 80 cm versus 40 cm rows index 42 percent compared to its monoculture yield
(Walker et al. 1984). Soybean in 20 cm rows yield- (Hashem et al. 2000).
ed more than in 40 cm or 80 cm rows when sickle- Grain yield of the rice cultivar Lemont was
pod was not controlled (Walker et al. 1984). Redroot reduced 21 percent when plants were within 25 cm
pigweed was 29 percent of total leaf area in wide (76 of a barnyardgrass plant group (four plants in 140
cm) rows and only 15 percent in narrow (25 cm) cm2 (Stauber et al. 1991). Rice yield was not affect-
rows (Lgre and Schreiber 1989). The leaf area dis- ed when the barnyardgrass plant group was 2550 or
tribution of soybean and redroot pigweed suggested 50100 cm from the rice. The optimum equidistant
light competition was important. With cultivation, rice plant spacing for optimal rice yield was 53 mm
38 cm rows resulted in less growth of redroot pig- to 71 mm (Counce et al. 1989). An increasing coef-
weed and robust foxtail than 76 cm rows, but with- ficient of variation in plant yield at closer spacings
out cultivation, the reverse was true (Orwick and is consistent with a large body of research that indi-
Schrieber 1979). cates that plant communities become more hierar-
In Ontario, Canada, the decrease in biomass pro- chical and variable as plant population densities
duction by transplanted and naturally occurring increase.
weeds was greater due to narrow row spacing than Final cotton emergence was not affected by cot-
to higher corn-population density. The combination tons planting pattern. At layby, there were more
of narrow rows (38 cm versus 76 cm) and high corn weeds when herbicide was not used in 51 cm com-
population increased corn canopy light interception pared to 102 cm rows, but at harvest the number of
3 to 5 percent (Begna et al. 2001). Weed biomass weeds was not different in the two row widths
was five to eight times lower under the corn canopy (Miller et al. 1983). There was no cotton yield
than in a weed monoculture. When corn density was advantage for narrow rows.
increased from seven to ten plants m-1 of row and With row spacings of 15, 25, 36, 46, and 91 cm, a
row width was decreased from 75 cm to 50 cm, there constant density of 43 snap beans m-2, and weed emer-
was significant increase in corns leaf area index and gence with the beans, only row spacings of 15 cm, 25
a reduction in the photosynthetic photon flux avail- cm, and 36 cm suppressed weed growth (18 percent)
able below the corn canopy (Murphy et al. 1996). In compared to standard 91 cm rows (Teasdale and Frank
all cases, narrow corn rows and increased corn- 1983). When weeds were controlled for the first half
population density significantly reduced the bio- of the growing season, 15 cm, 25 cm, and 36 cm rows
mass of late-emerging weeds. Corn yield increased suppressed weed growth 82 percent compared to 91-
10 to 15 percent in narrow (50-cm) rows, but intra- cm rows. Narrow rows suppressed weed growth by
specific corn competition in the higher density increasing the speed of canopy closure of snap bean
plantings significantly reduced early corn growth rows. Snap beans in 15 cm to 46 cm rows produced
and that offset the gain in yield from reduced weed similar yields that were higher by an average of 23
competition (Murphy et al. 1996). percent than the yield in 91 cm rows. With increasing
Decreasing wheat row spacing from 23 cm to 8 distance of horsenettle from snap bean rows, the
cm increased wheat yield at two of three locations weeds effect was reduced (Frank 1990).
and increased cheat infestation at six of ten locations Tomato density (0, 5, 10, or 20 plants m-1 of row)
(Koscelny et al. 1990). Increasing wheat planting and barnyardgrass density (0, 0.25, 0.5, 1, 2, 5, and
rate from 265 to 530 seeds m-2 increased wheat more than 50 plants m-1 of tomato crop row) had lit-
yield. tle effect on phenological development of barnyard-
Additional experiments in Oklahoma (Roberts et al. grass (Norris et al. 2001).
2001) showed that wheat row spacing did not affect With a constant in-row seeding rate, peanut yield
rye seed production. Averaged over all row spacings, increased as row width decreased from 81.2 cm (the
Weed Management Using the Principles of Competition 149

standard) to 40.6 cm and 20.3 cm. Weed growth was showed that in early season, weed interference
always less with narrower peanut rows. Without accounted more for yield reductions in monocultur-
weeds, peanut yields were 6 to 20 percent higher in al crops of maize and cowpea than it did in the
20.3 cm than in 81.2 cm rows (Buchanan and mixed culture of maize and cowpea.
Hauser 1980). Related work that has not focused on crop-weed
It is apparent, although not mentioned often, that competition has emphasized cover crops or living
the crops competitiveness increases with the percent- mulches that can be used as intercrops or companion
age of the field surface that it occupies. This percent- plants to suppress weeds. A review (Hartwig and
age is maximized as the crops planting pattern Ammon 2002) includes 93 references on the use of
achieves the greatest degree of rectangularity (the crop cover crops and living mulches for weed manage-
row spacing divided by the crops in-row spacing) ment. Hartwig and Ammon (2002) report that work
(Mohler 2001). In spite of the strong intuitive and the- with perennial living mulches such as crownvetch,
oretical basis (Fischer and Miles 1973 as cited by flatpea, birdsfoot trefoil, and white clover has shown
Mohler (2001) for an inverse relationship between that the living mulch does not have to be reseeded
crop row spacing and weed growth, Mohler (2001) annually. These plants, used as living mulches, con-
found only 27 of 49 studies, in 19 crops growing with serve nitrogen, reduce soil erosion, and increase soil
weeds, in which narrowing row spacing actually organic matter while reducing weed populations and
increased crop yield. The difference (Mohler 2001) losses due to weeds.
may be due to the effect of weed and crop height Appropriate weed control practices in farming
rather than just the area of ground covered. systems must consider the need to maintain soil fer-
tility and prevent erosion, and open row crops are
inimical to these needs. Akobundu (1980) developed
The applicable principles in polyculture and inter- integrated low or no-tillage weed management sys-
or companion cropping are: tems, compatible with more than one crop plant in a
field, which reduced herbicide use, fertilizer
1. Plant diversity is good and often diminishes
requirements, and soil erosion. Studies of a combi-
harmful interference.
nation of a legume or Eugusi melon and sweet pota-
2. Filling all ecological niches diminishes com-
to with corn, showed that these companion crops or
living mulches maintained corn yield, contributed to
3. A greater crop yield and less weed growth
nitrogen supply, suppressed weed growth, and
may be achieved if intercrops are more effec-
reduced soil erosion. In unweeded no-till plots, corn
tive than sole crops in usurping resources
grain yield was 1.6 T ha-1, whereas with convention-
from weeds or suppressing weed growth
al tillage it was 2.3 T ha-1. Corn yield in unweeded,
through allelopathy (Liebman and Dyck
live mulch plots averaged 2.7 T ha-1. Yields were not
different, and the presence of live mulch plants did
These principles have not been explored much in not reduce yield; they were complementary, not
weed science research. Weed scientists have, in a competitive.
very real sense, been bound by the dictum that the Clover has been grown successfully with corn and
only good plant in a field is the one that is planted has reduced weed growth (Vrabel et al. 1980). Crim-
and all others are to be regarded as weeds and, if son clover and subterranean clover were the most
possible, eliminated. It is part of weed sciences promising cover crops in cucumbers and peppers in
operative paradigm. Weed science research has Georgia and contributed to effective management of
made enormous progress toward achieving the goal diseases, nematodes, and insects (Phatak et al.
of clean monocultural fieldsfields without weeds. 1991). Sweet corn in a living mulch of white clover
The fact that research and the resultant technology had high yields in early years but lower yields later
have allowed this to be achieved in so many crops in because a contact herbicide used over the corn row
so many environments is laudable. On the other allowed invasion of perennial weeds that were not
hand, our paradigm has not urged us to explore the suppressed by white clover (Mohler 1991).
possibility that some plants may grow cooperatively Companion cropping (i.e., polyculture) can be a
or, at least, not competitively with crops. weed control technique, but research is needed to
Only one paper reviewed for this book was on determine how appropriate it may be in specific sit-
mixed or polyculture of plants. Ayeni et al. (1984a) uations. Limited evidence supports the contention
150 Chapter 8

that it can provide weed competition, build soil tard seeded at 530 seeds m-2 that grew with corn for
organic matter, reduce soil erosion, and improve four weeks. Weed growth in corn was reduced 51
water penetration (Andres and Clement 1984). percent and corn yield was reduced only 4 percent
When spring soil moisture is limiting, cover or com- compared to monocultural corn. DeHaan et al.
panion crops can deplete moisture and be detrimen- (1994) suggested it might be possible to develop
tal to crops in spite of weed control advantages. spring-seeded smother crops that reduce weed
In Pennsylvania, a polyculture of crownvetch, a growth up to 80 percent and have only a minor effect
legume, was successful as a living mulch in no- on corn. This innovative weed control technique has
tillage corn (Cardina and Hartwig 1980; Hartwig not been pursued.
1987). Crownvetch is difficult to establish, but once Another example of a weed used to gain interspe-
established it provides soil erosion control, cific competition in a polyculture is azolla for weed
improves fertility by reducing nutrient loss via ero- management in lowland rice. Azolla pinnata, a free-
sion, and contributes nitrogen and weed control. floating fern, has been tried in Asian rice culture
Weed control must be supplemented with herbicides because of its symbiotic relationship with Azolla
that will not kill the crownvetch. The system is anabena, a nitrogen-fixing blue-green algae. This
amenable to rotation of corn with other crops. symbiotic relationship can contribute up to 100 kg
Work in Ohio demonstrated use of hairy vetch for of nitrogen ha-1. A second use is for weed control
weed management. Unsuppressed hairy vetch due to the competitive effect of an azolla blanket
reduced weed biomass in corn 96 percent in one over the surface of paddy water. Perennial weeds
year and 58 percent in another. When corn was such as rushes and annuals with strong culms (e.g.,
planted in late April into hairy vetch in the early bud barnyardgrass) are not suppressed and must be con-
stage of growth, corn yield was reduced up to 76 trolled in other ways. Many other weeds are con-
percent. Hairy vetch competition was reduced or trolled well.
eliminated when corn was planted into hairy vetch Azolla has been successful but cannot be univer-
in mid- or late-bloom in May or early June. Because sally recommended because there is an increase in
of the shortened growing season and competition labor just to manage it. Some land must be devoted
from hairy vetch, corn planted in May into untreat- to supplying a continuing source of inoculum of
ed hairy vetch yielded similarly to corn planted in azolla for paddies and azolla may complicate other
the no-cover crop, weed-free check. pest problems. In fact, azolla may become a weed.
In Wisconsin, spring-planted winter rye has been These methods are not perfected and will not be
a successful living mulch for weed control in soy- the perfect answer to all weed problems. Polyculture
beans (Ateh and Doll 1996). A system employing is an incompletely explored weed management
just rye for weed control reduced weed shoot bio- opportunity. Such opportunities lead to lengthy
mass from 60 to 90 percent over three years. Rye research programs and are hard work. They chal-
worked best for weed control and did not reduce lenge the existing paradigm.
soybean yield when weed density was low and
ground cover from the mulch and soil moisture were
adequate for growth. Rye interference with soy- On arable land, tillage alone or in combination with
beans was minimal if rye was killed within 45 days other weed management methods may be an ade-
after soybean planting. quate system. Tillage turns under crop residue, con-
The concept of smother plants for weed control is ditions soil, and facilitates drainage. It controls
well known but not widely practiced. DeHaan et al. weeds by burying them, separating shoots from
1994) proposed the novel idea that it might be pos- roots, stimulating germination of dormant seeds and
sible to develop spring-seeded smother plants that buds (to be controlled by another tillage), desiccat-
reduce weed biomass early in the growing season, ing shoots, and depleting carbohydrate reserves of
but because they could be designed to live only for perennial weeds.
four to six weeks (the early critical weed control Other reasons for tillage include breaking up
time), they would have only a small or no effect on compacted soil, soil aeration, seed bed preparation,
corn yield. DeHaan et al. (1994) used yellow mus- trash incorporation, and crop cultivation. All of
tard selected to provide weed interference durations these are important, but the main accomplishment of
of 2, 4, 6, or 8 weeks and to grow only 10 cm to 20 most tillage done in the worlds crops is weed con-
cm tall. The best result was 10-cm-tall yellow mus- trol. The advent of no-till farming and minimum-till
Weed Management Using the Principles of Competition 151

farming have shown that tillage is not essential to from depth, differential survival at different depths,
grow crops and may do no more than control weeds. and the depth of seed burial with no-tillage, Mohler
Too frequent tillage can increase soil compaction, a (1993) made several predictions. First, in the first
disadvantage. Other disadvantages include exposure year following seed input to soil, nontilled areas will
of soil to erosion, moisture loss, and stimulation of have more emerged weed seedlings than tilled areas.
weed growth by encouraging germination of dor- In later years, no-till areas will have fewer emerged
mant seeds and vegetative buds. In some soils, with- weeds unless innate or induced dormancy is high. If
out tillage, soil can crust and there will be poor seed return is allowed, no-till areas will always have
water penetration. Decisions about the role of tillage more seedlings. After a major seed addition, plow-
must be made for each soil type and farming system. ing followed by years of shallow tillage is the best
Conclusions about tillages role, may be valid only management technique (Mohler 1993). A risk of no-
for the place and conditions of each study (Oryokot tillage systems is the development of perennial
et al. 1997). weeds. Over 14 years, a greater and more diverse
There are situations where plowing and subse- population of perennial weeds developed in reduced
quent tillage cannot prepare land for planting. These tillage systems than in moldboard plowed systems
include land heavily infested with perennial sod- (Buhler et al. 1994).
forming grasses, often encountered in developing Successful mechanical control of weeds is also
country agriculture. Many tillage implements give determined by human factors. Gunsolus (1990)
inadequate results in the crop row after the crop has noted that science could explain why certain weed
emerged and begun to grow. Tillage between rows is management practices work the way they do. Sci-
efficient. Crops can be cultivated to within a few ence develops basic principles to guide action.
inches of the row, but not as well in the crop row Human cultural knowledge is different from scien-
except by moving soil and burying weeds. To maxi- tific knowledge, although each may work toward the
mize tillage benefits, uniform spacing of crop rows, goal of good weed management. Cultural knowl-
straight rows achieved by precision planting, gauge edge tells one when and how to do something on a
wheels, and depth guides are needed. Uneven stands given soil and farm. Tillage is a cultural practice and
and driver error often lead to damage from mechan- therefore, by definition, it requires cultural knowl-
ical cultivation and destruction of some crop plants. edge. It requires the mind of a good farmer who
The success of tillage for weed control is deter- knows the land. Successful mechanical control
mined by biological factors: requires managerial skill (cultural knowledge) that
cannot be acquired from science. Such knowledge is
1. How closely weeds resemble the crop. Weeds acquired by doing and by observing those who have
that share a crops growth habit and time of done things well. Cultural knowledge is the art of
emergence may be the most difficult to con- farming whereby one knows how to select and apply
trol with tillage, especially when they grow in scientific knowledge to solve problems. Successful
crop rows. Weeds that emerge earlier or later mechanical control of weeds, regardless of imple-
than the crop are often easier to control. ment, is always related to the timeliness of the oper-
2. If a weeds seeds have a short, specific period ation. Research can determine when to do
of germination, it is easier to control them by something, but knowing when on a particular farm is
tillage as opposed to those whose seeds ger- part of the cultural knowledge good farmers have.
minate over a long time. There is no question that soil tillage and crop cul-
3. Perennial weeds that reproduce vegetatively tivation can control weeds and that tillage or
are particularly difficult to control with reduced tillage affect future weed populations. Sev-
tillage alone. eral studies have shown that reducing tillage affects
the population dynamics of annual weeds (Buhler
Tillages success is also determined by physical 1992; Buhler and Daniel 1988; Buhler and Oplinger
and environmental factors such as how wet soil is 1990; Johnson et al. 1989). The advent of no-tillage
and whether its condition prevents tillage. A wet practices to reduce soil disturbance and soil erosion
spring may prevent crop cultivation when weeds are have shown that tillage is not a required agricultural
small and controlled easily. Based on a model in practice for crop growth or weed control. However,
which the density of weed seedlings emerging is the data reveal that the effects of tillage are not con-
related to differing seedlings ability to emergence sistent among crops or across years and locations.
152 Chapter 8

Studies of the effects of tillage system on weeds cumulative seedling emergence in no-till plots. The
have not produced consistent results for all weeds. It effect was attributed to an extended dry period. The
is commonly observed that the effects of tillage, if presence or absence of corn did not affect common
present, are less important than the effects of crop and lambsquarters emergence or seedling density.
climate (Thomas and Frick 1993). Defelice et al. Tillage, as expected, reduced the weeds seedling
(1988) found no difference in the effect of conven- density, but the largest variation in seedling density
tional or no-tillage systems on control of velvetleaf in was attributed to varying environmental conditions
corn. In all three years of a study in Wisconsin that as is true for pigweeds (Oryokot et al. 1997).
compared the effect on weeds of moldboard plowing, The variable effects of different types of tillage on
chisel plowing, ridge tillage, and no-tillage, green different weed species are illustrated by work in
foxtail density was higher in chisel plowing and no- Nebraska that showed that ridge tillage enhanced
tillage than with moldboard plowing, and ridge tillage development of kochia and reduced density of wild
had the lowest density (Buhler 1992). Common proso millet and common lambsquarters. Tandem
lambsquarters density was always highest with chisel disking increased longspine sandbur and redroot
plowing (500 m-2 versus 75). Redroot pigweed aver- pigweed density, whereas moldboard plowing
aged 307 and 245 m-2 in no-till and chisel plow sys- increased common sunflower density (Wilson
tems versus only 25 in the other systems (Buhler 1993). It is probable that these effects would hold
1992). Oryokot et al. (1997) provide a reason for across years in this location, and they may hold for
these differences: pigweed seedlings emerge only these weeds in other locations, but these assump-
from the top 2.5 cm of soil regardless of tillage. Pig- tions must be tested.
weed seedling density is usually higher with no- The effect of four tillage systems (varying from
tillage because more seeds are nearer the soil surface. intensive to no tillage) on weed populations and ver-
Therefore, although tillage is necessary in many weed tical seed distribution was studied at three locations
management systems, it is less important than crop in Alberta (ODonovan and McAndrew 2000). The
and weather for pigweed population dynamics winter annual weeds, field pennycress, shepherds-
(Oryokot et al. 1997). Without weeds, corn yield was purse, and flixweed and the summer annuals wild
not affected by tillage system (Buhler 1992). Both the buckwheat and common lambsquarters all increased
tillage system and crop rotation altered the relation- in the soil seedbank as tillage increased. Thus, the
ship between corn yield over two years but tillage was effect of increased tillage intensity on common
not a factor in soybean yield in one year (McGiffen et lambsquarters was consistent across locations. In the
al. 1997). In Nigeria, with minimum or no weed inter- Alberta study, increased soil seedbank populations
ference, corn yield was better with conventional did not always result in increased weed seedling
tillage (plowing followed by two harrowings) than populations with no tillage, which ODonovan and
no-tillage, but it was worse when weeds were present McAndrew (2000) suggested may mean that the
(Ayeni et al. 1984b). requirement for herbicidal weed control may be
Bararpour and Oliver (1998) found that with reduced with no-tillage systems. In contrast, and
tillage 11 percent of the soil seedbank of common to illustrate the difficulty of extrapolating these
cocklebur and sicklepod emerged one year after the results across locations and weed species, both soil
seed fell on the soil, but with no-tillage only 0.7 per- seedbank and spring weed seedling populations of
cent of common cocklebur and 1.6 percent of sick- shepherds-purse at two locations and of flixweed at
lepod seed emerged the next year. With tillage, another were highest in the zero tillage system. In
common cocklebur became the dominant weed, but contrast, the soil seedbank and spring seedling pop-
with no-tillage, sicklepod dominated. The seedbank ulation of green foxtail decreased as tillage
of both species was depleted faster with tillage. decreased, suggesting that it should become less of
The effects of three tillage systems (no-till, chisel a problem as tillage decreases. The effects of tillage
plow, and moldboard plow) and the presence or are confounded by crop residues. In Wisconsin, over
absence of corn on soil temperature, moisture, and three years with varied environmental conditions,
the emergence and density of common lambsquar- when tillage affected giant foxtail and redroot fox-
ters were studied at two sites in Ontario, Canada tail, emergence was greater in untilled than tilled
(Roman et al. 1999). Tillage system affected the (simulated moldboard plowing by spading 20 cm
phenology of the weeds emergence in only one year deep) plots (Buhler et al. 1996). Velvetleaf emer-
when more days were required to reach 80 percent gence was greater from tilled than untilled soil in
Weed Management Using the Principles of Competition 153

two of three years, and the effects on common tillage system (moldboard plow, chisel plow, and
lambsquarters were not consistent over three years. no-till), whereas redroot pigweed density was usual-
Maize surface residue was inconsistent on giant fox- ly higher in the chisel plow system. Moldboard
tail and common lambsquarters. Velvetleaf emer- plowing always had greater velvetleaf density than
gence was reduced by two or four times the base no-till, and the latter always had greater giant foxtail
level of residue, and the effect of maize residue on density. Giant foxtail and redroot pigweed became
redroot pigweed emergence was dependent on more difficult to control when tillage was reduced,
tillage and precipitation (Buhler et al. 1997). Reduc- but velvetleaf became easier to control (Buhler and
ing tillage has a greater effect on the population Oplinger 1990). Another example is work in Ontario
dynamics of the four annual weeds than surface that showed that the response of annual dicots and
maize residues (Buhler et al. 1997). monocots to tillage was inconsistent in oats, barley,
Esbenshade et al. (2001) found that burcucumber and wheat (Lgre and Bai 1999). Perennial dicots
emergence frequency was independent of tillage dominated in no-till systems in the three small
system (no-till versus reduced tillage). Preplant grains, whereas perennial monocots were more
tillage increased the number of emerged burcucum- abundant in tilled systems in all three cereals
ber plants by 110 percent in one year and 70 percent (Lgre and Bai 1999). One must also conclude that
in another compared to no tillage. Johnsongrass pro- the effects of tillage are not consistent between
duces longer rhizomes with limited tillage. These crops. For example, oat and barley populations were
rhizomes, as opposed to those broken by tillage, will not affected by no-tillage but wheat population was
grow more rapidly and johnsongrass interference reduced 16 to 20 percent in no-till systems (Lgre
will begin earlier as tillage decreases (Lolas and and Bai 1999). The effects of tillage may be consis-
Coble 1980). tent for a weed species across locations.
A study of cultivation frequency and time of initi- A final point about the effects of tillage concerns
ation showed that seed cotton yield was increased at sampling to measure the effect. Mulugeta et al.
three of nine locations when cultivation was initiat- (2001) point out that the relationship between
ed two weeks instead of one week after emergence species richness and sample area has been shown in
(Colvin et al. 1992). Cultivation with a flexible tine many natural communities but has rarely been con-
harrow in the fall reduced density of common chick- sidered in crop-weed communities. Using sampling
weed, catchweed bedstraw, and rape, and thinned areas ranging from 0.0625 to 512 m2 in 14 nested
but did not reduce the yield of wheat. Yield was sample areas, they determined the influence of sam-
maintained because 1000 grain weight increased but ple area on species richness. The functional mini-
the number of grains did not (Wilson et al. 1993). mum area required to represent 75 percent of the
Summer biomass of common chickweed and catch- total weed species in tilled and short-term no-till
weed bedstraw was reduced more by spring than by fields was 32 m2. No functional minimum area was
fall harrowing, but biomass of rape was reduced determined in long-term, no-till fields because
only by fall harrowing. Wilson et al. (1993) con- species richness continued to increase over the range
cluded that weakly rooted, climbing or decumbent of sample areas. Regression analysis indicated that
species are more easily controlled by spring cultiva- sample areas of less than 1 m2 would contain less
tion whereas species that develop a tap root are more than 50 percent of the observed maximum species
readily controlled by tillage at an early growth stage richness in a field. Sample areas of 36 m2 in tilled
in the fall. It is highly probable that this conclusion and short-term, no-till fields and 185 m2 in long-
may be applicable across locations and species. term, no-till fields would measure 75 percent of the
Mulugeta and Boerboom (2000) showed that observed maximum species richness. Therefore, as
there was variation in the onset of the critical time of chapter 9 proposes, how one does the work is a
weed removal in soybeans between a reduced tillage major determinant of what one can conclude from
and no-tillage system between years and within what was done. It is as important in determining the
tillage systems across years. Therefore, based on effect of tillage on weed species as it is in all other
several studies, one must conclude that tillage has an areas of weed management decision making.
effect but it is not consistent among weed species.
For example, in Wisconsin, a two-year study (Buh-
ler and Oplinger 1990) showed that common lambs- The first edition of this book included only five
quarters density was not greatly influenced by papers that reported effects of crop rotation on weed
154 Chapter 8

management (Zimdahl 1980). In spite of evidence of never occur in rice. Nightshades are common in pota-
the utility of crop rotation for weed management, lit- toes, tomatoes, and beans, and kochia and lambs-
tle additional research has been done since 1979. quarters are frequent in sugarbeets. Dandelions are
The literature survey by Liebman and Dyck (1993a) common in turf but not as common in row crops,
found that weed population density and weed bio- although without management, dandelions can
mass were reduced by crop rotation (what they increase in row crops and in pastures and hay (e.g.,
called temporal diversification) and intercropping alfalfa).
(spatial diversification). Compared to monoculture, These associations occur because of similarity in
crop rotation reduced weed density in 21 cases, crop and weed phenology (naturally occurring phe-
increased it in only 1 case, and made no difference nomena that recur periodically, e.g., flowering),
in 5 cases. Twelve studies reported effects on weed- adaptation to cultural practices (e.g., tillage, mow-
seed density, which was lower in nine and equal in ing, irrigation), similar growth habits (e.g., time to
three studies. Liebman and Dyck (1993a) report that mature or to reach full height), and perhaps of most
the success of crop rotation for weed management is importance, resistance or adaptation to imposed
based on varying patterns of resource competition, weed control methods. When one crop is grown for
allelopathy, soil disturbance, and mechanical dam- many years (monoculture), weeds, present in the
age to seedlings that create an unstable and fre- soil seedbank, will be favored and their populations
quently inhospitable environment that prevents the will increase. Weed-crop associations are not acci-
proliferation of a particular weed species. It is clear dental and can be explained. Associations can be
from Liebman and Dycks (1993a) review that the changed by changing crop, time of planting, or weed
effect of crop rotation on weeds is well known and control method. Wild oats can be reduced in small
supported theoretically, but data in support of the grain crops by growing corn in the rotation and
theory are lacking. Most of the evidence in support using herbicides selective in corn plus cultivation for
of the benefits and wisdom of crop rotation for weed control when corn is grown. The same practices can-
management is observational and anecdotal rather not be used when small grain crops are grown.
than the result of carefully planned studies that sys- A good rotation includes crops that reduce (man-
tematically manipulate specific components of rota- age) weeds that are especially troublesome in suc-
tional systems to isolate and improve those elements ceeding crops. Management is accomplished by
(e.g., interrow cultivation, choice of crop genotype) competition or through use of different weed control
or combinations of elements that may be important techniques in different crops. In many places, barley
for weed control (Liebman and Dyck 1993a). The is planted in spring before soil temperatures are
weed management effects of crop rotations, while ideal for germination of most weeds. An exception
generally accepted, should be assessed through is common lambsquarters, which can be a serious
careful study of extant, complex farming systems weed in barley. Beans, on the other hand, are plant-
and the design and testing of new integrated ed in late spring after tillage has destroyed many, but
approaches (Liebman and Dyck 1993a). It seems not all, summer annual weeds.
that although the benefits of crop rotation are Ball and Miller (1990) showed that weed species
accepted by farmers and researchers, there is a composition varied with cropping sequence among
paucity of research data to support the benefits in rotations of corn for three years, pinto beans for
modern agricultural systems. Rotation regularly three years, or two years of sugarbeets followed by
changes the crop, soil preparation practices, subse- one year of corn. Hairy nightshades seedbank pop-
quent soil tillage, and weed control techniques in a ulation increased after three years of pinto beans;
field. All of these affect weed populations, and while green foxtail increased after three years of corn; and
crops are not commonly rotated to control weeds, the sugarbeet-corn sequence caused an increase in
the effect of rotation as a determinant of weed prob- kochia. Ball and Miller (1990) attributed the differ-
lems is accepted. ences to the herbicides used in each cropping
If it is done, crops are rotated for economic, mar- sequence. Crop cultivation, land preparation time
ket, and agronomic reasons but rarely for weed man- and method, and time of planting and harvest may
agement. It is known that some weeds associate with also favor one weed and discourage others.
certain crops more than with others. Barnyardgrass Two weeds dominated the relative dry weight of
and junglerice are common in rice. Wild oats are weeds in four cropping systems in the Philippines,
common in irrigated wheat and barley but almost but their relative magnitude in the cropping systems,
Weed Management Using the Principles of Competition 155

on the same soil, was different (Pablico and Moody difficult to control in winter wheat. Downy brome
1984). In a rice-sorghum rotation, itchgrass domi- density increased from 24 to 970 plants m-2 over five
nated, but with continuous sorghum, itchgrass near- years in Alberta, Canada, and density was often
ly disappeared and spiny amaranth dominated. higher with no tillage (Blackshaw 1994). When fal-
Different cropping systems affect weed populations low or spring canola were rotated with winter wheat,
and may favor or deter species. downy brome population was reduced to less than
Growing competitive crops (e.g., hemp) in rota- 55 or 100 plants m-2, respectively, over six years.
tion may complement other means of control of yel- Blackshaw (1994) concluded that continuous winter
low nutsedge, which is sensitive to competition for wheat cropping is not a good option in areas where
light. Growth and reproduction (tuber production downy brome is prevalent, and either a fallow/wheat
and density in a following corn crop) of yellow or fallow/canola/wheat rotation is better.
nutsedge in the Netherlands was reduced in corn The importance of rotation for management of
that followed corn grown for silage, winter rye sicklepod was emphasized by Johnson et al. (1994).
grown for silage, winter barley, and hemp compared Sicklepod growing alone in fallowed areas produced
to corn following on land with no preceding crop more seed per plant and more seedlings than when
(Lotz et al. 1991). After hemp, hardly any yellow the weed grew with a crop. Sicklepod growing in
nutsedge tuber production was observed. Growing a corn produced fewer seeds per plant than when it
green manure crop after barley harvest reduced yel- grew with peanuts or cotton.
low nutsedge tuber production to 40 percent of that Long-term studies to determine the effect of dif-
in winter barley followed by fallow. Competition for ferent cropping sequences on the population dynam-
light was the main reason for reduction in yellow ics of winter wild oat (Fernandez-Quintanilla et al.
nutsedge growth and reproduction. 1984) showed that continuous winter cereal crop-
When corn was intercropped with cassava, corn ping (with or without herbicides) increased the win-
yield decreased with time in weeded and unweeded ter wild oat soil seedbank from 26 to 80 percent per
plots except in corn plots followed by one to three year. With spring barley, the soil seedbank declined
years of weeded Pueraria fallow. Similarly, cassava 10 percent per year. When sunflower was a summer
tuber yield decreased with time in all unweeded plots crop or a 12-month fallow was included in the rota-
in all treatments except when cassava followed a tion to prevent new seed production, the soil seed
weeded Pueraria fallow (Akobundu et al. 1999). The reserve declined 57 to 80 percent annually. There
results of this work in Nigeria suggest that rotations was a great reduction in the size of the soil seedbank
that included fallow years during which the soil was of winter wild oats if the cropping program was
planted with a legume species offered more effective other than continuous winter cereals (Fernandez-
weed management than natural bush fallow. Quintanilla et al. 1984).
Work on intercropping of pea and barley with the
weed white mustard showed that nitrogen supply,
water supply, soil conditions, and pea genotype The role of shade in reducing plant vigor and growth
(height) all interacted (Liebman 1989). To fully is well known and exploited in weed management
evaluate intercropping, the desired yield of the c