The Weedy Rice Problem
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FAO Weedy rices – origin, PLANT PRODUCTION biology, ecology AND PROTECTION and control PAPER 188 by James C. Delouche Nilda R. Burgos David R. Gealy Gonzalo Zorrilla de San Martín and Ricardo Labrada with the collaboration of Michael Larinde and Cadmo Rosell FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS Rome, 2007 5IFNFOUJPOPSPNJTTJPOPGTQFDJm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ª'"0 iii Contents Acknowledgements x Preface xi List of acronyms xiii 1. Introduction 1 2. The weedy rice problem 3 What weedy rices are 3 Where weedy rices are a problem 5 The origins and sources of weedy rices 9 Why weedy rices are so successful as weeds 13 3. Diversity of weedy rice populations 17 Early and general observations on the diversity and variability of red rice 18 Louisiana – an early study of variability in red rice 19 Mississippi – a case study of diversity and variability in red rice 20 Texas – comparative study of diversity in red rices from four states 28 Arkansas – phenotypic and genetic diversity of red rice 30 Hybrid swarms 34 Hybridization of red and cultivated rice in the United States of America 37 4. Seed shattering and dormancy in weedy rices 45 Early and heavy seed shattering 45 Intense and persistent seed dormancy 51 5. Vigour and competitiveness of weedy rices 63 Economic losses from weedy rices 63 Competitive traits of red rices compared with cultivated rices 64 Effects of red rice density and period of interference 67 Overall effects of competition 70 The physiology and competitiveness of red rices in the United States of America 70 6. Ecological relationships 79 Seed shattering and dormancy 79 Variation in maturity 81 Protracted emergence 82 Environmental interactions on and in the soil 85 Ecological model of red rice germination 92 Further discussion on hybrid swarms from cultivated rice / red rice crosses 93 iv 7. Strategies for controlling weedy rices 97 Recognition of weedy rices 97 Preventing infestations 99 Depletion of the weedy rice soil seed bank 104 Suppression of weedy rice germination (water management and chemicals) 106 Destruction or removal of weedy rice plants in the rice crop 108 Alternation of rice with other crops or field uses to change environment 109 Biotechnological strategy – herbicide-resistant varieties 111 Requisites for effective weedy rice control programmes 113 8. Examples of weedy rice control campaigns 115 South America – combat and control of red rice in Uruguay 115 Central America and the Caribbean – management and control of weedy rices 121 North America 124 9. Conclusions 127 References 131 v List of tables 1. Major characteristics of the spikelets and grains in 1 084 panicles 19 2. Characteristics of 42 red rice contaminants from commercial rice fields in Mississippi and Texas, USA, 1978–1990 as compared with four cultivated varieties 22 3. Mean values for three traits influencing competitiveness 24 4. Characteristics of progeny from the 78/21 red rice segregate collected in a field of Starbonnet rice in Mississippi, USA, in 1978 27 5. Characteristics of progeny from the 79/16 red rice segregate collected in a field of the Starbonnet rice in Mississippi, USA, in 1979 29 6. Characterization of red rice in Arkansas, USA 33 7. Outcrossing from rice to rice and from rice to red rice under conditions in the United States of America 39 8. Estimated outcrossing rates between Clearfield rice (CL161) and different biotypes of red rice in commercial fields, Arkansas, USA 41 9. Shattering time and sees moisture content of 14 rice phenotypes 46 10. Time-course of seed shattering in selected red rice phenotypes and moisture content of seed in the upper third of the panicle at the first evidence of shattering 47 11. Time and degree of shattering of five weedy rice lines compared with variety IR 64 48 12. Intensity of seed dormancy for cultivated varieties and RR phenotypes harvested at 24 days past anthesis and dried for 9 days at 22–24 °C and after 6 months storage 53 13. Initial level of dormancy in 54 randomly selected SHR segregates 54 14. Variation in the persistence of dormancy in seeds of two cultivated varieties and five red rice phenotypes in open storage at 30 °C 54 15. Effects of storage of fully imbibed non-dormant and dormant seeds (circa 28-percent moisture content) of two red rice types at 30 °C on dormancy and germination 55 16. Differential germinative responses of non-dormant and dormant seeds of red rice ecotypes after accelerating ageing for periods up to 24 days 56 17. Effects of low temperatures and period of exposure on germination responses of fully imbibed non-dormant and dormant seeds (27–30-percent moisture content) of the Nato variety and the BLKH and SHA+ red rice phenotypes 56 18. Temperature ranges for germination for cultivated varieties and red rice ecotypes at intervals during a 14-day period 66 19. Comparison of seedling vigour for the Maybelle and Lemont varieties and the SHA- and BLKH red rice phenotypes during an 8-day period at 25 °C in dark conditions 67 20. Effects of season-long competition between the Starbonnet variety and the SHA- and BLKH red rice types in a 1:1 ratio on the time of 50-percent anthesis, harvest maturity, tiller number, plant dry weight, and grain yield 68 vi 21. Effects of season-long competition between the Maybelle and Lemont varieties and the SHA- and BLKH red rice types in transplant and direct seeded culture on biomass and some yield components of the two varieties 69 22. Comparative agronomic and physiological traits of rice and weedy rice grown in non-competitive conditions at the Rice Research and Extension Center, Stuttgart, Arkansas, between May and August in 2000 and 2001 72 23. Comparative growth between rice and red rice under non-competitive condition, with optimal soil fertility level, in pot experiments conducted outdoors, May–September 2001, at Main Agricultural Research and Extension Center, University of Arkansas, Fayettev 76 24. Cumulative emergence percentages over an 8-month period of three cultivated varieties and four red rice types planted on 28 September 1981 82 25. Effects of planting depth for two different dates on emergence and seedling height of the Starbonnet variety and the SHA-, SHA+ and BLKH red rices 20 and 30 days after planting 84 26. Germination and dormancy of seeds after air storage in the field and buried 15 cm deep in soil under rainfed and flooded conditions 87 27. Germination and dormancy percentages of dormant and non-dormant seeds after periods of submergence 20 cm deep in water under ambient conditions 89 28. Effects of periods of constant and alternating temperatures on germination and dormancy of seeds of the SHA+ red rice ecotype 90 29. Effects of storage temperature and seed moisture content on the induction of dormancy in seeds of the SHA+ red rice phenotype 91 30. Effect of temperature regime in releasing induced dormancy in seeds of the SHA+ red rice phenotype 92 31. Seed testing laboratory standards for red rice in Uruguay 117 32. Survey of analyses of incoming rice received by the main rice mills of Uruguay 119 vii List of figures 1. Red rice distribution by hull colour in commercial fields in Arkansas, United States of America, summer 2002 and 2003 10 2. Rice production areas in the United States of America, 2005 18 3. Multidimensional scale diagram (using GD values derived from RAPD markers) of genotypic clustering of red rice accessions and cultivated rice 32 4. Effects of storage temperature on the rate of dormancy release in seeds of the Nato variety and five red rice phenotypes 59 5. Mean lengths of the mesocotyl, coleoptile, and exserted plumule for two rice varieties and two red rice phenotypes after 10 days at 25 °C 66 6. Mean lengths of the mesocotyl, coleoptile, and exserted plumule for two rice varieties and two red rice phenotypes after 8 days at 30 °C 67 7. Relative shoot dry weights of rice cultivar and red rice ecotypes at 70 days after emergence 74 8. Comparative response of Drew rice and strawhull red rice to fertilizer N in terms of biomass production and N-use efficiency, two weeks after heading 75 9. Cumulative field emergence over a 15-month period of the Starbonnet variety and three red rice phenotypes planted after 0, 2, 4 and 6-week simulated delays in shattering in 1983 80 10. Cumulative emergence over a 13-month period of seeds of the Labelle variety and three red rice phenotypes planted on 15 September 1982 83 11. Cumulative emergence over a 13-month period of seeds of the Labelle variety and three red rice phenotypes planted on 9 June 1983 83 12. Seasonal variation of dormancy and germination in deeply buried seeds of the Nato variety and three red rices 88 13. Evolution of the rice area in Uruguay, 1930–2005 115 14. Evolution of the total planted area and the area planted with certified seeds in Uruguay, 1988–1998 117 15. National system for the production of rice seed in Uruguay 118 viii List of plates 1. Typical strawhull-red (SHR) and blackhull-red (BHR) rices collected in Mississippi in 1978 and 1979. Note lax and loose panicle branches and spikelets (seeds) lined up like beads on the branches. Seeds are shown in inserted photos. 6 2.