Herbicides are the foundation of weed control in commercial crop-production systems. However, herbicide-resistant (HR) weed populations are evolving rapidly as a natural response to selection pressure imposed by modern agricultural management activities. Mitigating the evolution of herbicide resistance depends on reducing selection through diversification of weed control techniques, minimizing the spread of resistance genes and genotypes via pollen or propagule dispersal, and eliminating additions of weed seed to the soil seedbank. Effective deployment of such a multifaceted approach will require shifting from the current concept of basing weed management on single-year economic thresholds.

The herbicide glyphosate became widely used in the United States and other parts of the world after the commercialization of glyphosate-resistant crops. These crops have constitutive overexpression of a glyphosate-insensitive form of the herbicide target site gene, 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS). Increased use of glyphosate over multiple years imposes selective genetic pressure on weed populations. We investigated recently discovered glyphosate-resistant Amaranthus palmeri populations from Georgia, in comparison with normally sensitive populations. EPSPS enzyme activity from resistant and susceptible plants was equally inhibited by glyphosate, which led us to use quantitative PCR to measure relative copy numbers of the EPSPS gene. Genomes of resistant plants contained from 5-fold to more than 160-fold more copies of the EPSPS gene than did genomes of susceptible plants. Quantitative RT-PCR on cDNA revealed that EPSPS expression was positively correlated with genomic EPSPS relative copy number. Immunoblot analyses showed that increased EPSPS protein level also correlated with EPSPS genomic copy number. EPSPS gene amplification was heritable, correlated with resistance in pseudo-F 2 populations, and is proposed to be the molecular basis of glyphosate resistance. FISH revealed that EPSPS genes were present on every chromosome and, therefore, gene amplification was likely not caused by unequal chromosome crossing over. This occurrence of gene amplification as an herbicide resistance mechanism in a naturally occurring weed population is particularly significant because it could threaten the sustainable use of glyphosate-resistant crop technology.5-enolpyruvylshikimate-3-phosphate synthase | herbicide resistance | mobile genetic element | evolution | Palmer amaranth

A glyphosate-resistant Palmer amaranth biotype was confirmed in central Georgia. In the field, glyphosate applied to 5- to 13-cm-tall Palmer amaranth at three times the normal use rate of 0.84 kg ae ha−1controlled this biotype only 17%. The biotype was controlled 82% by glyphosate at 12 times the normal use rate. In the greenhouse,I50values (rate necessary for 50% inhibition) for visual control and shoot fresh weight, expressed as percentage of the nontreated, were 8 and 6.2 times greater, respectively, with the resistant biotype compared with a known glyphosate-susceptible biotype. Glyphosate absorption and translocation and the number of chromosomes did not differ between biotypes. Shikimate was detected in leaf tissue of the susceptible biotype treated with glyphosate but not in the resistant biotype.

In little over 20 yr, Palmer amaranth has risen from relative obscurity to its current status as one of the most widespread, troublesome, and economically damaging agronomic weeds in the southeastern U.S. Numerous factors have enabled Palmer amaranth to become such a dominant and difficult-to-control weed, including its rapid growth rate, high fecundity, genetic diversity, ability to tolerate adverse conditions, and its facility for evolving herbicide resistance. It is both a serious threat to several U.S. cropping systems and a fascinating model weed. In this paper, we review the growing body of literature on Palmer amaranth to summarize the current state of knowledge on the biology, agricultural impacts, and management of this weed, and we suggest future directions for research.

Development of herbicide-resistant crops has resulted in significant changes to agronomic practices, one of which is the adoption of effective, simple, low-risk, crop-production systems with less dependency on tillage and lower energy requirements. Overall, the changes have had a positive environmental effect by reducing soil erosion, the fuel use for tillage, and the number of herbicides with groundwater advisories as well as a slight reduction in the overall environmental impact quotient of herbicide use. However, herbicides exert a high selection pressure on weed populations, and density and diversity of weed communities change over time in response to herbicides and other control practices imposed on them. Repeated and intensive use of herbicides with the same mechanisms of action (MOA; the mechanism in the plant that the herbicide detrimentally affects so that the plant succumbs to the herbicide; e.g., inhibition of an enzyme that is vital to plant growth or the inability of a plant to metabolize the herbicide before it has done damage) can rapidly select for shifts to tolerant, difficult-to-control weeds and the evolution of herbicide-resistant weeds, especially in the absence of the concurrent use of herbicides with different mechanisms of action or the use of mechanical or cultural practices or both.

Changes in the weed flora of cropping systems reflect the impacts of factors that create safe sites for weed establishment and facilitate the influx and losses to and from the soil seedbank. This analysis of the annual surveys of the Southern Weed Science Society documents changes in the weed flora of the 14 contiguous southern states since the advent of transgenic, herbicide-resistant crops. In 1994 and 2009, the top five weeds in corn were morningglories, Texas millet, broadleaf signalgrass, johnsongrass, and sicklepod; in this same period Palmer amaranth, smartweeds, and goosegrass had the greatest increases in importance in corn. In cotton, morningglories and nutsedges were among the top five most troublesome weeds in 1995 and 2009. Palmer amaranth, pigweeds, and Florida pusley were also among the five most troublesome species in 2009; the weeds with the largest increases in importance in cotton were common ragweed and two species with tolerance to glyphosate, Benghal dayflower and Florida pusley. In soybean, morningglories, nutsedges, and sicklepod were among the top five weed species in 1995 and 2009. Two species with glyphosate resistance, Palmer amaranth and horseweed, were the second and fourth most troublesome weeds of soybean in 2009. In wheat, the top four weeds in 2008 were the same as those in 1994 and included Italian ryegrass, wild garlic, wild radish, and henbit. Crop production in the southern region is a mosaic of various crop rotations, soil types, and types of tillage. During the interval between the surveys, the predominant change in weed management practices in the region and the nation was the onset and rapid dominance of the use of glyphosate in herbicide-resistant cultivars of corn, cotton, and soybean. Because of the correspondence between the effects of glyphosate on the respective weed species and the observed changes in the weed flora of the crops, it is likely the very broad use of glyphosate was a key component shaping the changes in weed flora. Only eight of the top 15 most troublesome weeds of cotton and soybean, the crops with the greatest use of glyphosate, were the same in 1995 and 2009. In contrast, in corn and wheat where adoption of glyphosate-resistant cultivars lags or is absent, 12 of the 15 most troublesome weeds were the same in 1994 and 2008. These findings show on a regional scale that weeds adapt to recurrent selection from herbicides, currently the predominant weed management tool. Future research should seek methods to hinder the rapid spread of herbicide-tolerant and evolution of herbicide-resistant weed species. As new tools are developed, research should focus on ways to preserve the efficacy of those tools through improved stewardship.

The Southern Weed Science Society has conducted an annual survey of the most troublesome weeds in several major crops since 1971. The objective of this summary was to characterize shifts in weed populations over a 22-yr period in four major agronomic crops. For corn, soybean, cotton, and peanut, the largest increases in rank as the most troublesome weeds were found with sicklepod and bermudagrass. The largest decreases were found with johnsongrass, crab-grasses, and common cocklebur. Morningglories and nutsedges remained relatively constant weed problems over the 22-yr period. Sicklepod, nutsedges, and morningglories were the three most troublesome weeds averaged over all crops because they are so well established and relatively difficult to control. Pigweeds (Palmer amaranth, sandhills amaranth, tumble pigweed, and water-hemps) have become increasingly important in soybean, peanut, and cotton in a limited number of states.

Organic systems in the southeastern USA offer unique challenges and solutions to crop production due to regional soil and climate characterized by highly weathered soil types, high precipitation and the capacity to grow cover crops in the winter. Recently, the interest of producers and researchers in high-residue cover crops and conservation tillage systems has increased. Various designs of the roller-crimper to manage cover crops have been invented and demonstrated to growers in the southeastern region of the USA over the past 17 years. The impacts of high-residue cover crop mulches on the agronomic systems in the region are diverse. Legume cover crops assist with meeting N demand from cash crops though they decompose rapidly and are seldom sufficient for N demanding crops such as corn. Cereal cover crop mulches can have the opposite effect by immobilizing N and have a longer impact on soil moisture and weed dynamics. While undesirable for many crops, N immobilization is one possible mechanism for weed suppression in legume cash crops planted into cereal residues. Other cover crop weed suppression mechanisms include physical impedance, light availability, allelopathy and microclimate effects. Regardless of the cause, successful weed control by mulches is highly dependent on having substantial biomass. The southeastern region is capable of producing cover crop biomass in excess of 9000 kg ha − 1 , which is sufficient for weed control in many cash crops, although supplementary weed control is sometimes necessary. Long-term data are needed to predict when farmers should add supplementary weed control. More work is also needed on how much additional N is required for the cash crops and how best to deliver that N in a high-residue environment using organic sources.