No abstract

In input-intensive cropping systems around the world, farmers rarely proactively manage weeds to prevent or delay the selection for herbicide resistance. Farmers usually increase the adoption of integrated weed management practices only after herbicide resistance has evolved, although herbicides continue to be the dominant method of weed control. Intergroup herbicide resistance in various weed species has been the main impetus for changes in management practices and adoption of cropping systems that reduce selection for resistance. The effectiveness and adoption of herbicide and nonherbicide tactics and practices for the proactive and reactive management of herbicide-resistant (HR) weeds are reviewed. Herbicide tactics include sequences and rotations, mixtures, application rates, site-specific application, and use of HR crops. Nonherbicide weed-management practices or nonselective herbicides applied preplant or in crop, integrated with less-frequent selective herbicide use in diversified cropping systems, have mitigated the evolution, spread, and economic impact of HR weeds.

The frequency of gene flow from Brassica napus L. (canola) to four wild relatives, Brassica rapa L., Raphanus raphanistrum L., Sinapis arvensis L. and Erucastrum gallicum (Willd.) O.E. Schulz, was assessed in greenhouse and/or field experiments, and actual rates measured in commercial fields in Canada. Various marker systems were used to detect hybrid individuals: herbicide resistance traits (HR), green fluorescent protein marker (GFP), species-specific amplified fragment length polymorphisms (AFLPs) and ploidy level. Hybridization between B. rapa and B. napus occurred in two field experiments (frequency approximately 7%) and in wild populations in commercial fields (approximately 13.6%). The higher frequency in commercial fields was most likely due to greater distance between B. rapa plants. All F(1) hybrids were morphologically similar to B. rapa, had B. napus- and B. rapa-specific AFLP markers and were triploid (AAC, 2n=29 chromosomes). They had reduced pollen viability (about 55%) and segregated for both self-incompatible and self-compatible individuals (the latter being a B. napus trait). In contrast, gene flow between R. raphanistrum and B. napus was very rare. A single R. raphanistrum x B. napus F1 hybrid was detected in 32,821 seedlings from the HR B. napus field experiment. The hybrid was morphologically similar to R. raphanistrum except for the presence of valves, a B. napus trait, in the distorted seed pods. It had a genomic structure consistent with the fusion of an unreduced gamete of R. raphanistrum and a reduced gamete of B. napus (RrRrAC, 2n=37), both B. napus- and R. raphanistrum-specific AFLP markers, and had <1% pollen viability. No hybrids were detected in the greenhouse experiments (1,534 seedlings), the GFP field experiment (4,059 seedlings) or in commercial fields in Québec and Alberta (22,114 seedlings). No S. arvensis or E. gallicum x B. napus hybrids were detected (42,828 and 21,841 seedlings, respectively) from commercial fields in Saskatchewan. These findings suggest that the probability of gene flow from transgenic B. napus to R. raphanistrum, S. arvensis or E. gallicum is very low (<2-5 x 10(-5)). However, transgenes can disperse in the environment via wild B. rapa in eastern Canada and possibly via commercial B. rapa volunteers in western Canada.

Diagnosing herbicide-resistant weeds as a first step in resistance management and monitoring their nature, distribution, and abundance demands efficient and effective screening tests. This review summarizes and recommends appropriate seed sampling techniques, protocols for screening weeds for resistance to herbicides of different sites of action, interpretation of results, and information given to the grower. Elements common to all screening procedures are reviewed. Choosing appropriate discriminating doses to distinguish between resistant and susceptible weed biotypes is the most important factor in achieving accurate and consistent results. Interpretation of results is also critical because resistant weeds may comprise a small portion of the population in suspected accessions or biotypes.

Herbicide rotations and mixtures are widely recommended to manage herbicide resistance. However, little research has quantified how these practices actually affect the selection of herbicide resistance in weeds. A 4-yr experiment was conducted in western Canada from 2004 to 2007 to examine the impact of herbicide rotation and mixture in selecting for acetolactate synthase (ALS) inhibitor resistance in the annual broadleaf weed, field pennycress, co-occurring in wheat. Treatments consisted of the ALS-inhibitor herbicide, ethametsulfuron, applied in a mixture with bromoxynil/MCPA formulated herbicide (photosystem-II inhibitor/synthetic auxin), or in rotation with the non-ALS inhibitor at an ALS-inhibitor application frequency of 0, 25, 50, 75, and 100% (i.e., zero to four applications, respectively) over the 4-yr period. The field pennycress seed bank at the start of the experiment contained 5% ethametsulfuron-resistant seed. Although weed control was only marginally reduced, resistance frequency of progeny of survivors increased markedly after one ALS-inhibitor application. At the end of the experiment, the level of resistance in the seed bank was buffered by susceptible seed, increasing from 29% of recruited seedlings after one application to 85% after four applications of the ALS inhibitor. The level of resistance in the seed bank for the mixture treatment after 4 yr remained similar to that of the nontreated (weedy) control or 0% ALS-inhibitor rotation frequency treatment. The results of this study demonstrate how rapidly ALS-inhibitor resistance can evolve as a consequence of repeated application of herbicides with this site of action, and supports epidemiological information from farmer questionnaire surveys and modeling simulations that mixtures are more effective than rotations in mitigating resistance evolution through herbicide selection.

Kochia [Kochia scoparia (L.) Schrad.] is an annual broadleaf weed species native to Eurasia and introduced as an ornamental to the Americas by immigrants in the mid- to late 1800s. Although sometimes categorized in the genus Bassia, there is no compelling reason for this classification. This naturalized species is a common and economically important weed in crop production systems and ruderal areas in semiarid to arid regions of North America, and has expanded northward in the Canadian Prairies during the past 30 yr. Although primarily self-pollinated, substantial pollen-mediated gene flow and efficient seed dispersal aids both short- and long-distance spread. The weed is morphologically highly variable, and its growth and development are markedly affected by environmental conditions. Kochia, a C4 species, is highly competitive in cropping systems because of its ability to germinate at low soil temperatures and emerge early, grow rapidly, tolerate heat, drought and salinity, and exert allelopathic effects on neighboring species. Moreover, herbicidal control has been compromised to some extent by the widespread evolution of herbicide resistance in the species. Kochia is used as a forage, is palatable to livestock with nutritional value similar to that of alfalfa (Medicago sativa), but can be toxic if it comprises the majority of the diet. Although kochia pollen is an allergen, the seed is a source of phytochemicals including mosquito pheromones and saponins that are potentially beneficial to human health; kochia also is beneficial in phytoremediation of soils contaminated by hydrocarbons or pesticides. Key words: Kochia, Kochia scoparia, Bassia scoparia, herbicide resistance, soil salinity tolerance, weed biology

Multiple herbicide resistance to glyphosate, glufosinate, bromoxynil, or imidazolinone in volunteer plants of canola (Brassica napus) has been attributed to pollen flow among cultivars with different resistance traits. A study was conducted in Saskatchewan, Canada, in 1999 and to assess gene flow in space and time in adjacent commercial fields of glyphosate-and glufosinate-resistant canola, including (1) estimation of gene flow with distance; (2) frequency and distribution of volunteers, and effect on gene flow; (3) effect of adventitious double herbicide-resistant seed presence in seedlots planted; and (4) a comparison of various marker systems to track gene flow events. At 11 sites in 1999, gene flow was determined by sampling seeds from plants located at 0, 50, 100, 200, 400, 600, or 800 m along a transect perpendicular to the common border in the paired fields, spraying seedlings with glyphosate and glufosinate, and confirming the presence of the transgenes using commercial test strips and PCR analysis. In the spring of 2000, putative double herbicide-resistant volunteers that survived sequential herbicide applications were mapped at three of the sites using GPS and resistance in sampled plants was characterized. In 1999, gene flow between the paired fields was detected to a maximum distance of 400 m. Values ranged from 1.4% outcrossing at the border common to the paired fields to 0.04% at 400 m. In 2000, gene flow as a result of pollen flow in 1999 was detected to the limits of the study areas (800 m). Large variation in gene flow levels and patterns among the three sites was evident. Adventitious presence of double herbicide-resistant seed in glyphosate-resistant seedlots planted at two of the sites in 1999 contributed to the occurrence of double herbicide-resistant volunteers in 2000. The results of this study suggest that gene stacking in B. napus canola volunteers in western Canada may be common, and reflects pollen flow between different herbicide-resistant canola, presence of double herbicideresistant off-types in seedlots, and/or agronomic practices typically employed by Canadian growers.

This review is both a retrospective (what have we missed?) and prospective (where are we going?) examination of weed control and technology, particularly as it applies to herbicide-resistant weed management (RWM). Major obstacles to RWM are discussed, including lack of diversity in weed management, unwillingness of many weed researchers to conduct real integrated weed management research or growers to accept recommendations, influence or role of agrichemical marketing and governmental policy and lack of multidisciplinary research. We then look ahead to new technologies that are needed for future weed control in general and RWM in particular, in areas such as non-chemical and chemical weed management, novel herbicides, site-specific weed management, drones for monitoring large areas, wider application of 'omics' and simulation model development. Finally, we discuss implementation strategies for integrated weed management to achieve RWM, development of RWM for developing countries, a new classification of herbicides based on mode of metabolism to facilitate greater stewardship and greater global exchange of information to focus efforts on areas that maximize progress in weed control and RWM. There is little doubt that new or emerging technologies will provide novel tools for RMW in the future, but will they arrive in time?