Carol A. Mallory-Smith

Revised Classification of Herbicides by Site of Action for Weed Resistance Management Strategies1

The classification of herbicides by site of action, published in 1997, has been revised. The classification system uses a numbering system for a herbicides site of action, chemical family, and common name. Regulatory agencies in the United States and Canada have published labeling guidelines based on the classification to aid in herbicide resistance management. Abbreviations: EPA, Environmental Protection Agency; HRAC, Herbicide Resistance Action Committee; PMRA, Pest Management Regulatory Agency.

Herbicide Resistance: Toward an Understanding of Resistance Development and the Impact of Herbicide-Resistant Crops

This is the publisher’s final pdf. The published article is copyrighted by the Weed Science Society of America and can be found at: http://wssajournals.org/loi/wees. To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work.

Establishment of transgenic herbicide‐resistant creeping bentgrass (Agrostis stolonifera L.) in nonagronomic habitats

Concerns about genetically modified (GM) crops include transgene flow to compatible wild species and unintended ecological consequences of potential transgene introgression. However, there has been little empirical documentation of establishment and distribution of transgenic plants in wild populations. We present herein the first evidence for escape of transgenes into wild plant populations within the USA; glyphosate‐resistant creeping bentgrass (Agrostis stolonifera L.) plants expressing CP4 EPSPS transgenes were found outside of cultivation area in central Oregon. Resident populations of three compatible Agrostis species were sampled in nonagronomic habitats outside the Oregon Department of Agriculture control area designated for test production of glyphosate‐resistant creeping bentgrass. CP4 EPSPS protein and the corresponding transgene were found in nine A. stolonifera plants screened from 20 400 samples (0.04 ± 0.01% SE). CP4 EPSPS‐positive plants were located predominantly in mesic habitats downwind and up to 3.8 km beyond the control area perimeter; two plants were found within the USDA Crooked River National Grassland. Spatial distribution and parentage of transgenic plants (as confirmed by analyses of nuclear ITS and chloroplast matK gene trees) suggest that establishment resulted from both pollen‐mediated intraspecific hybridizations and from crop seed dispersal. These results demonstrate that transgene flow from short‐term production can result in establishment of transgenic plants at multi‐kilometre distances from GM source fields or plants. Selective pressure from direct application or drift of glyphosate herbicide could enhance introgression of CP4 EPSPS transgenes and additional establishment. Obligatory outcrossing and vegetative spread could further contribute to persistence of CP4 EPSPS transgenes in wild Agrostis populations, both in the presence or absence of herbicide selection.

Gene flow from glyphosate-resistant crops.

Gene flow from transgenic glyphosate-resistant crops can result in the adventitious presence of the transgene, which may negatively impact markets. Gene flow can also produce glyphosate-resistant plants that may interfere with weed management systems. The objective of this article is to review the gene flow literature as it pertains to glyphosate-resistant crops. Gene flow is a natural phenomenon not unique to transgenic crops and can occur via pollen, seed and, in some cases, vegetative propagules. Gene flow via pollen can occur in all crops, even those that are considered to be self-pollinated, because all have low levels of outcrossing. Gene flow via seed or vegetative propagules occurs when they are moved naturally or by humans during crop production and commercialization. There are many factors that influence gene flow; therefore, it is difficult to prevent or predict. Gene flow via pollen and seed from glyphosate-resistant canola and creeping bentgrass fields has been documented. The adventitious presence of the transgene responsible for glyphosate resistance has been found in commercial seed lots of canola, corn and soybeans. In general, the glyphosate-resistant trait is not considered to provide an ecological advantage. However, regulators should consider the examples of gene flow from glyphosate-resistant crops when formulating rules for the release of crops with traits that could negatively impact the environment or human health.

Glyphosate-Resistant Italian Ryegrass (Lolium multiflorum) in California: Distribution, Response to Glyphosate, and Molecular Evidence for an Altered Target Enzyme

Abstract Selection by herbicides has resulted in widespread evolution of herbicide resistance in agricultural weeds. In California, resistance to glyphosate was first confirmed in rigid ryegrass in 1998. Objectives of this study were to determine the current distribution and level of glyphosate resistance in Italian ryegrass, and to assess whether resistance could be due to an altered target site. Seeds were sampled from 118 populations and seedlings were treated with glyphosate at 866 g ae ha−1. Percentage of survivors ranged from 5 to 95% in 54 populations. All plants from 64 populations died. One susceptible (S) population, four putatively resistant (R) populations, and one S accession from Oregon were used for pot dose–response experiments, shikimic acid analyses, and DNA sequencing. Seedlings were treated with glyphosate at eight rates, ranging from 108 to 13,856 g ae ha−1. Shoot biomass was evaluated 3 wk after treatment and fit to a log-logistic regression equation. On the basis of GR50 (herbicide rate required to reduce growth by 50%) values, seedlings from putatively R populations were roughly two to 15 times more resistant to glyphosate than S plants. Shikimic acid accumulation was similar in all plants before glyphosate treatment, but at 4 and 7 DAT, S plants from California and Oregon accumulated approximately two and three times more shikimic acid, respectively, than R plants. Sequencing of a cDNA fragment of the EPSPS coding region revealed two different codons, both of which encode proline at amino acid position 106 in S individuals. In contrast, all R plants sequenced exhibited missense mutations at site 106. Plants from one population revealed a mutation resulting in a proline to serine substitution. Plants from three R populations exhibited a mutation corresponding to replacement of proline with alanine. Our results indicate that glyphosate resistance is widespread in Italian ryegrass populations of California, and that resistance is likely due to an altered target enzyme. Nomenclature: Glyphosate; Italian ryegrass, Lolium multiflorum Lam. LOLMU; rigid ryegrass, Lolium rigidum Gaud. LOLRI.

Identification of glyphosate-resistant Italian ryegrass (Lolium multiflorum) in Oregon

Abstract A suspected glyphosate-resistant Italian ryegrass biotype was collected from a filbert orchard near Portland, OR, where glyphosate was applied multiple times per year for about 15 yr. Greenhouse studies were conducted to determine if this biotype was glyphosate resistant. The plants were sprayed with glyphosate (0.01 to 3.37 kg ae ha−1) 14 d after planting and shoot biomass was determined 3 wk after herbicide treatment. Based on the dose–response experiments conducted in the greenhouse, the suspected Italian ryegrass biotype was approximately fivefold more resistant to glyphosate than the susceptible biotype. Plants from both susceptible and resistant biotypes were treated with glyphosate (0.42 and 0.84 kg ha−1) and shikimic acid was extracted 12, 24, 48, and 96 h after treatment. The susceptible biotype accumulated between three and five times more shikimic acid than did the resistant biotype. Leaf segments from both susceptible and resistant biotypes were incubated with different glyphosate concentrations (0.5 to 3000 μM) for 14 h under continuous light. Shikimic acid was extracted from each leaf segment and quantified. At a concentration up to 100 μM, leaf segments from the susceptible biotype accumulated more shikimic acid than leaf segments from the resistant biotype. The epsps gene was amplified and sequenced in both susceptible and resistant biotypes; however, no amino acid change was found in the resistant biotype. The level of resistance in this biotype is similar to that reported for a glyphosate-resistant Italian ryegrass biotype from Chile. Nomenclature: Glyphosate; Italian ryegrass, Lolium multiflorum Lam. LOLMU; filbert, Corylus avellana L.

The fertility of wheat × jointed goatgrass hybrid and its backcross progenies

Abstract The spontaneous flow of genes from wheat to jointed goatgrass is of great concern to breeders intending to release herbicide-resistant wheat. The objectives of this research were to study how genes could flow from wheat to jointed goatgrass through crossing and backcrossing between these two species and, based on this knowledge, to propose possible ways to minimize the chance of gene flow between them. Results showed that the wheat × jointed goatgrass hybrid can only serve as a female parent to produce the BC1 generation. The BC1 generation was found to have 1.8% male fertility and 4.4% female fertility, indicating that it could serve as either the male or female parent to produce a BC2 generation. The fertility of the resultant BC2 generation further increased. The average male, female, and self-fertility was 8.9, 18.0, and 6.9%, respectively. After the BC2 generation, the backcross progeny has three possible ways to reproduce: to pollinate jointed goatgrass, to be pollinated by jointed goatgrass, or to pollinate itself. Restoration of the chromosome number of jointed goatgrass continues as the BC2 generation is selfed, but some plants can contain an alien chromosome over generations. The possible ways to reduce the chance of gene flow between these two species are (1) prevent the production of hybrids, (2) prevent the production of the BC1 generation, and (3) put a herbicide-resistant gene on the A- or B-genome of wheat. Nomenclature:Jointed goatgrass; Aegilops cylindrica Host AEGCY; wheat; Triticum aestivum L.

Evolution of Weediness and Invasiveness: Charting the Course for Weed Genomics

Abstract The genetic basis of weedy and invasive traits and their evolution remain poorly understood, but genomic approaches offer tremendous promise for elucidating these important features of weed biology. However, the genomic tools and resources available for weed research are currently meager compared with those available for many crops. Because genomic methodologies are becoming increasingly accessible and less expensive, the time is ripe for weed scientists to incorporate these methods into their research programs. One example is next-generation sequencing technology, which has the advantage of enhancing the sequencing output from the transcriptome of a weedy plant at a reduced cost. Successful implementation of these approaches will require collaborative efforts that focus resources on common goals and bring together expertise in weed science, molecular biology, plant physiology, and bioinformatics. We outline how these large-scale genomic programs can aid both our understanding of the biology of weedy and invasive plants and our success at managing these species in agriculture. The judicious selection of species for developing weed genomics programs is needed, and we offer up choices, but no Arabidopsis-like model species exists in the world of weeds. We outline the roadmap for creating a powerful synergy of weed science and genomics, given well-placed effort and resources.

Seed production on Triticum aestivum by Aegilops cylindrica hybrids in the field

Abstract Field experiments were conducted to determine if seeds would be produced on Triticum aestivum by Aegilops cylindrica hybrids in the field and, if it were, to determine the viability of the seeds produced. One, five, or 10 hybrids were planted into varying proportions of A. cylindrica and T. aestivum in a replacement series design. Percent seed set ranged from 0 to 5.5% in 1996 and from 0 to 9.2% in 1997. Seeds were set in all treatments. The average seed set was 2.3% in 1996 and 3.8% in 1997. No differences in seed set were found among treatments. The seeds produced were separated according to seed condition, either full or shriveled, and tested for germination. The germination of the seeds produced on the hybrids was not significantly different between years. The average germination for full seeds was 94% in both years and 79 and 84% for shriveled seeds in 1995 and 1996, respectively. Greenhouse studies were conducted to evaluate the rate of self-fertility of the BC1 generation and to identify morphological characteristics that could be used to identify the probable pollen donor parent and to predict the occurrence of seed set. In 1997 4.1% and in 1998 2.1% of BC1 plants set seeds. The average seed set was 0.3% in 1997 and 0.06% in 1998. It was not possible, using any morphological characteristic measured, to determine the identity of the parent serving as the pollen donor in the previous generation or to predict the occurrence of seed set in the BC1 generation. This is the first reported study to show that hybrids between T. aestivum and A. cylindrica have the ability, although limited, to backcross under field conditions and set seeds. Furthermore, the seeds produced are viable and will germinate and produce plants. With the millions of hectares of T. aestivum infested with A. cylindrica, even the limited ability to backcross is of concern for the movement of a herbicide-resistance gene. Nomenclature: Aegilops cylindrica Host AEGCY, jointed goatgrass; Triticum aestivum L., wheat.

SULFONYLUREA HERBICIDE RESISTANT WEEDS: DISCOVERY, DISTRIBUTION, BIOLOGY, MECHANISM, AND MANAGEMENT

Sulfonylurea herbicide resistant Lactuca serriola L. (prickly lettuce) plants were discovered near Lewiston, Idaho in April 1987. This was the first confirmed occurrence of herbicide resistance resulting from the use of sulfonylurea. Kochia scoparia (L.) Schrad (kochia) plants from Liberal, Kansas were confirmed as resistant to chlorsulfuron and metsulfuron-methyl in 1988. Since then, sulfonylurea resistant K. scoparia has been identified in eight other States and one Canadian Province. Salsola iberica Sennen & Pau (russian thistle) plants from Kansas, Montana, North Dakota and Washington and Stellaria media (L.) Vill. (common chickweed) plants from Alberta, Canada have also been found to be resistant to sulfonylurea herbicides. Most resistant plants were collected from fields where dryland winter wheat (Triticum aestivum L.) was grown either continuously or in rotation with summer fallow and where chlorsulfuron or chlorsulfuron plus metsulfuron-methyl had been applied at 7 to 14 month intervals for 3 to 5 years. Total sulfonylurea herbicide used ranged from 52 to 200 g a.i.ha-1. Resistant K. scoparia plants have also been collected from noncrop areas where sulfometuron-methyl was applied annually for 3 to 4 years. Total sulfometuron-methyl used ranged from 96 to 425 g ai ha-1. The mechanism of resistance is an altered site of action, acetohydroxyacid synthase enzyme, which is inhibited less in resistant than in susceptible biotypes by sulfonylurea and imidazolinone herbicides. Resistance is not due to differences in herbicide absorption, translocation, or metabolism. Chlorsulfuron-resistant K. scoparia and L. serriola are resistant to several other sulfonylurea and imidazolinone herbicides. However, resistant and susceptible biotypes are usually controlled equally by herbicides with different modes of action. Resistance in L. serriola is controlled by one nuclear gene with incomplete dominance.