In Defense of Crop Monoculture

Because yields typically decline, crop monoculture is commonly considered as not sustainable. This yield decline is due largely to soilborne plant pathogens adapted to/specialized for the roots of the crop. For high-value fruit and vegetable crops, yields are maintained with monoculture using soil fumigation or soil solarization. Soils can also be sanitized by flooding, which may account for the success of paddy rice monocultures. Our work in the U.S. Pacific Northwest has focused on four root and crown diseases of wheat and barley, namely take-all caused by Gaeumannomyces graminis var. tritici, Fusarium crown rot caused by Fusarium culmorum and Fusarium pseudograminearum, Rhizoctonia root rot caused by Rhizoctonia solani AG8 and R. oryzae, and Pythium root rot caused by several Pythium species. Herein, we describe a remarkable and apparently wide-spread microbiological control (disease suppression) in the rhizosphere that is responsible for the well-documented decline of take-all and coordinate increase in crop yield following one or more outbreaks of the disease and continued monoculture of wheat or barley. Since this disease suppression is specific for take-all, other strategies are under development for control of the other three root and crown diseases with wheat-intensive cropping systems, including in direct-seed systems. The strategies include the development of transgenic resistance in barley to Rhizoctonia root rot using the ThEn42 gene from Trichoderma harzianum for production of a 42-Kda endochitinase, selection of wheat cultivars for tolerance to Fusarium crown rot, and use of a systemic seed-treatment chemicals and current-year seed for seedling protection against Pythium root rot. Introduction Crop rotation, which we define as the practice of growing different crops in sequence in the same field, is nearly as old as agriculture itself and remains the centerpiece of most cropping systems worldwide. Crop monoculture, which we define as the practice of replanting the same crop species in the same field, with no break to a different crop, has an equally long but less successful history, except for some high-value fruit and vegetable crops grown with the aid of soil fumigation or soil solarization, and also possibly for paddy rice. “Crop monoculture” is also used to describe large areas planted to the same crop species, e.g. the vast area planted to wheat each year in the North American Great Plains. For purposes of our defense of crop monoculture, like crop rotation, we mean crop monoculture in the temporal and not the spatial sense. Growers that practice crop monoculture generally do so for economic reasons. The selected crop is the most profitable and any profitability loss from yield declines are less than that which occurs from any rotational options available. In these situations, the ability to minimize the losses associated with monoculture can provide the best option to increase productivity and profitability. Many names have been given to the phenomenon of yield decline with monoculture, including replant problem, autotoxicty, and monoculture injury. Even perennial crops such as alfalfa and grasses are prone to show yield (or stand) decline over time, typically starting in the third or fourth year following their establishment. Similarly with annual crops, yields typically decline starting in the third or fourth year of the monoculture, although some yield decline may occur already in the second year of monoculture (Cook and Baker, 1983). Because of these yield declines, crop monoculture is commonly considered as not sustainable. Crop rotation, like tillage, is an invention of agriculture. Indeed, annual plants in the wild reseed themselves in more or less the same places year after year—and without tillage. The occurrence of wild plants in polycultures could provide a kind of rotational benefit in cases where the seeds of one species happen to fall on the site occupied the previous year by a different species. In the case of wheat, the

[1]  M. Gerlagh Introduction of Ophiobolus graminis into new polders and its decline , 1968, Netherlands Journal of Plant Pathology.

[2]  B. Landa,et al.  Interactions Between Strains of 2,4-Diacetylphloroglucinol-Producing Pseudomonas fluorescens in the Rhizosphere of Wheat. , 2003, Phytopathology.

[3]  R. Cook Take-all of wheat , 2003 .

[4]  J. T. de Souza,et al.  Frequency, Diversity, and Activity of 2,4-Diacetylphloroglucinol-Producing Fluorescent Pseudomonas spp. in Dutch Take-all Decline Soils. , 2003, Phytopathology.

[5]  B. Landa,et al.  Differential Ability of Genotypes of 2,4-Diacetylphloroglucinol-Producing Pseudomonas fluorescens Strains To Colonize the Roots of Pea Plants , 2002, Applied and Environmental Microbiology.

[6]  B. M. Gardener,et al.  Microbial populations responsible for specific soil suppressiveness to plant pathogens. , 2002, Annual review of phytopathology.

[7]  K. Sivasithamparam,et al.  Influence of depth of soil disturbance on root growth dynamics of wheat seedlings associated with Rhizoctonia solani AG-8 disease severity in sandy and loamy sand soils of Western Australia , 2001 .

[8]  D. Weller,et al.  Exploiting Genotypic Diversity of 2,4-Diacetylphloroglucinol-Producing Pseudomonas spp.: Characterization of Superior Root-Colonizing P. fluorescensStrain Q8r1-96 , 2001, Applied and Environmental Microbiology.

[9]  P. Butterworth,et al.  Genetic and pathogenic variation among cereal, medic and sub-clover isolates of Pythium irregulare , 2001 .

[10]  L. Thomashow,et al.  Genetic Diversity of phlD from 2,4-Diacetylphloroglucinol-Producing Fluorescent Pseudomonas spp. , 2001, Phytopathology.

[11]  B. Ownley,et al.  Influence of paired-row spacing and fertilizer placement on yield and root diseases of direct-seeded wheat. , 2000 .

[12]  S. Kalloger,et al.  Genotypic and Phenotypic Diversity of phlD-ContainingPseudomonas Strains Isolated from the Rhizosphere of Wheat , 2000, Applied and Environmental Microbiology.

[13]  D. Weller,et al.  Effect of Population Density of Pseudomonas fluorescens on Production of 2,4-Diacetylphloroglucinol in the Rhizosphere of Wheat. , 1999, Phytopathology.

[14]  L. Thomashow,et al.  Identification and Characterization of a Gene Cluster for Synthesis of the Polyketide Antibiotic 2,4-Diacetylphloroglucinol from Pseudomonas fluorescens Q2-87 , 1999, Journal of bacteriology.

[15]  C. Lawrence,et al.  Genes from mycoparasitic fungi as a source for improving plant resistance to fungal pathogens. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[16]  J. Browse,et al.  A role for jasmonate in pathogen defense of Arabidopsis. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[17]  A. Boronin,et al.  A Seven-Gene Locus for Synthesis of Phenazine-1-Carboxylic Acid by Pseudomonas fluorescens2-79 , 1998, Journal of bacteriology.

[18]  D. Weller,et al.  Natural plant protection by 2,4-diacetylphloroglucinol-producing Pseudomonas spp. in take-all decline soils , 1998 .

[19]  R. Cook,et al.  Bacillus sp. L324-92 for Biological Control of Three Root Diseases of Wheat Grown with Reduced Tillage. , 1997, Phytopathology.

[20]  L. Thomashow,et al.  Frequency of Antibiotic-Producing Pseudomonas spp. in Natural Environments , 1997, Applied and environmental microbiology.

[21]  L. Thomashow,et al.  Quantification of 2,4-Diacetylphloroglucinol Produced by Fluorescent Pseudomonas spp. In Vitro and in the Rhizosphere of Wheat , 1997, Applied and environmental microbiology.

[22]  K. K. Thomsen,et al.  Transgenic barley expressing a protein-engineered, thermostable (1,3-1,4)-beta-glucanase during germination. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[23]  C. Keel,et al.  Conservation of the 2,4-diacetylphloroglucinol biosynthesis locus among fluorescent Pseudomonas strains from diverse geographic locations , 1996, Applied and environmental microbiology.

[24]  A. Rovira,et al.  Effect of sowing point design and tillage practice on the incidence of Rhizoctonia root rot, take-all and cereal cyst nematode in wheat and barley , 1996 .

[25]  P. E. Rasmussen,et al.  Diseases of wheat in long-term agronomic experiments at Pendleton, Oregon. , 1996 .

[26]  A. Jagendorf,et al.  Molecular mechanisms of defense by rhizobacteria against root disease. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[27]  D. Weller,et al.  Purification of an antibiotic effective against Gaeumannomyces graminis var. tritici produced by a biocontrol agent, Pseudomonas aureofaciens , 1993 .

[28]  D. Backhouse,et al.  Long-term effects of stubble management on the incidence of infection of wheat by Fusarium graminearum Schw. Group 1 , 1993 .

[29]  R. D. Prew Wheat health management , 1992 .

[30]  L. Thomashow,et al.  Cloning and heterologous expression of the phenazine biosynthetic locus from Pseudomonas aureofaciens 30-84. , 1992, Molecular plant-microbe interactions : MPMI.

[31]  R. Cook Wheat root health management and environmental concern , 1992 .

[32]  D. Weller,et al.  Genetic analysis of the antifungal activity of a soilborne Pseudomonas aureofaciens strain , 1991, Applied and environmental microbiology.

[33]  L. Thomashow,et al.  Production of the Antibiotic Phenazine-1-Carboxylic Acid by Fluorescent Pseudomonas Species in the Rhizosphere of Wheat , 1990, Applied and environmental microbiology.

[34]  S. Neate A comparison of controlled environment and field trials for detection of resistance in cereal cultivars to root rot caused by Rhizoctonia solani , 1989 .

[35]  K. Sivasithamparam,et al.  Growth promotion of rotation crop species by a sterile fungus from wheat and effect of soil temperature and water potential on its suppression of take-all , 1989 .

[36]  K. Sivasithamparam,et al.  Pathogen-suppression: A case study in biological suppression of Gaeumannomyces graminis var. Tritici in soil , 1989 .

[37]  L. Thomashow,et al.  Role of a phenazine antibiotic from Pseudomonas fluorescens in biological control of Gaeumannomyces graminis var. tritici , 1988, Journal of bacteriology.

[38]  P. E. Rasmussen,et al.  Long-term Tillage and Nitrogen Fertilization Effects on Organic Nitrogen and Carbon in a Semiarid Soil , 1988 .

[39]  R. Allmaras,et al.  Effects of Soil Compaction and Incorporated Crop Residue on Root Health , 1988 .

[40]  R. Cook,et al.  Influence of soil treatments on growth and yield of wheat and implications for control of pythium root rot , 1987 .

[41]  C. Rothrock Take-all of wheat as affected by tillage and wheat-soybean doublecropping , 1987 .

[42]  R. Cook,et al.  Infection of wheat embryos by Pythium species during seed germination and the influence of seed age and soil matric potential , 1987 .

[43]  A. Rovira,et al.  Reduction of Rhizoctonia root rot of direct-drilled wheat by short-term chemical fallow , 1987 .

[44]  R. Cook,et al.  Application of a rapid screening test for selection of bacteria suppressive to take-all of wheat , 1985 .

[45]  R. Cook,et al.  Increased take-all of wheat with direct drilling in the Pacific Northwest , 1984 .

[46]  K. F. Baker,et al.  The nature and practice of biological control of plant pathogens , 1985 .

[47]  R. Cook The influence of rotation crops on take-all decline phenomenon. , 1981 .

[48]  R. Cook Fusarium foot rot of wheat and its control in the Pacific Northwest. , 1980 .

[49]  S. Wilhelm,et al.  How Soil Fumigation Benefits the California Strawberry Industry , 1980, Plant Disease.

[50]  R. Smiley Wheat-rhizoplane pseudomonads as antagonists of Gaeumannomyces graminis , 1979 .

[51]  A. Rovira Studies on soil fumigation—I: Effects on ammonium, nitrate and phosphate in soil and on the growth, nutrition and yield of wheat , 1976 .

[52]  A. Rovira,et al.  The role of bacteria in the biological control of Gaeumannomyces graminis by suppressive soils , 1976 .

[53]  J. Deacon Biological control of the take-all fungus, Gaeumannomyces graminis, by Phialophora radicicola and similar fungi , 1976 .

[54]  E. Ridge Studies on soil fumigation—II: Effects on bacteria , 1976 .

[55]  R. Papendick Plant Water Stress and Development of Fusarium Foot Rot in Wheat Subjected to Different Cultural Practices , 1974 .