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Weed-crop Competition: A Review PDF

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Weed-Crop Competition A Review Second Edition Weed-Crop Competition A Review Second Edition Robert L. Zimdahl Robert L. Zimdahlis Professor of Weed Science, Authorization to photocopy items for internal or per- Colorado State University,and edited the Journal sonal use,or the internal or personal use of specific Weed Sciencefrom 1994 to 2002. clients,is granted by Blackwell Publishing,provided that the base fee of $.10 per copy is paid directly to ©2004 Blackwell Publishing the Copyright Clearance Center,222 Rosewood Drive, All rights reserved Danvers,MA 01923. For those organizations that have been granted a photocopy license by CCC,a Blackwell Publishing Professional separate system of payments has been arranged. The 2121 State Avenue,Ames,Iowa 50014,USA fee code for users of the Transactional Reporting Ser- vice is 0-8138-0279-2/2004 $.10. Orders: 1-800-862-6657 Office: 1-515-292-0140 Printed on acid-free paper in the United States of Fax: 1-515-292-3348 America Web site: www.blackwellprofessional.com Library of Congress Cataloging-in-Publication Data Blackwell Publishing Ltd 9600 Garsington Road,Oxford OX4 2DQ,UK Zimdahl,Robert L. Tel.:+44 (0)1865 776868 Weed-crop competition :a review / Robert L. Zimdahl—2nd ed. Blackwell Publishing Asia p. cm. 550 Swanston Street,Carlton,Victoria 3053,Australia Includes bibliographical references (p.). Tel.:+61 (0)3 8359 1011 ISBN 0-8138-0279-2 (alk. paper) 1. Weeds. 2. Plant competition. 3. Weeds— Bibliography. 4. Plant competition—Bibliography. I. Title. SB611.Z55 2004 632'.5—dc22 2003023656 The last digit is the print number:9 8 7 6 5 4 3 2 1 In order to penetrate ever further into their subjects, the host of specialists narrow their fields and dig down deeper and deeper till they can’t see each other from hole to hole. But the treasures their toil brings to light they place on the ground above. A different kind of specialist should be sitting there, the only one still missing. He would not go down any hole, but would stay on top and piece all the different facts together. —Thor Heyerdahl,Aku-Aku:The Secret of Easter Island Contents Preface ix 1 Introduction:An Historical Perspective 1 2 Definition of Plant Competition 6 3 Competition in the Community 9 Plant Communities 9 Agricultural Communities 10 Levels of Competition 10 Density 13 Community Composition 14 Theories of Competition 16 4 Influence of Competition on the Plant 19 Density 19 Competitive Ability 19 General Principles 20 Competitive Success 22 5 The Effect of Weed Density 27 Alfalfa 28 Barley 28 Corn/Maize 32 Cotton 39 Oilseed Crops 46 Flax 46 Rapeseed/Canola 46 Safflower 48 Sunflower 48 Peanut/Groundnut 49 Potato 51 Rice 52 Sorghum 56 Soybean 57 Sugarbeet 75 Sugarcane 77 Vegetables 78 Bean 78 Lentil and Chickpea 80 vii viii Contents Onion 81 Pea 81 Pepper 82 Tomato 83 Other Vegetable Crops 86 Wheat 88 Other Small Grain Crops 103 Studies of Diverse Crops 104 Weed-Weed Interference 106 6 The Effect of Competition Duration 109 7 The Elements of Competition 131 The Role of Temperature 132 Competitive Interactions for Nutrients 133 Competitive Interactions for Light 136 Competitive Interactions for Water 139 Competition for Other Environmental Factors 141 8 Weed Management Using the Principles of Competition 146 Plant Arrangement in the Community 147 Monoculture Versus Polyculture 149 Tillage 150 Rotation or Crop Sequence 153 Shade 155 The Role of Crop Genotype 156 Fertility 159 The Importance of Weed Biology and Ecology 160 9 Methods Used to Study Weed-Crop Competition 167 10 Models and Modeling 173 Conceptual Models 174 Simulation (Analytical) Models 174 Mechanistic or Empirical Models 176 Time of Emergence 179 Leaf Area Models 179 Multispecies Competition 182 The Extrapolation Domain of Models 183 Decision-Aid Models 183 Spatial Distribution 185 The Effect of Variability on Decisions 186 Thresholds 187 Conclusion 190 11 Conclusion:The Complexity of Competition 197 Appendix 200 Index 211 Preface The primary impetus for the first edition of this 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. one’s 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 science—“the 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 Kuhn’s 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 IRRI’s 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 ix 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 book’s 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- Press. 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. Roget’s 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. Allelopathy—An update. Bot. Rev. Amer. Chem. Soc. Symp. Series No. 268. 45:15–109. Washington,DC. ———. 1983. Pest control with nature’s 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 1 Introduction: 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 3:17–18). 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 earth’s 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 vice. with respect to each other. He derived a theory of antagonism between phanerogams, and a theory of Malthus’s views are claimed by some to have crop rotation based on the idea that succeeding been inspiration for Darwin’s (1859) principle of species should be those not inhibited by toxic sub- natural selection. Darwin (1859) derived the concept 1

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