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The objective of this paper was to review the reproductive biology, herbicide-resistant (HR) biotypes, pollen-mediated gene flow (PMGF), and potential for transfer of alleles from HR to herbicide-susceptible grass weeds including barnyardgrass, creeping bentgrass, Italian ryegrass, johnsongrass, rigid (annual) ryegrass, and wild oats. The widespread occurrence of HR grass weeds is at least partly due to PMGF, particularly in obligate outcrossing species such as rigid ryegrass. Creeping bentgrass, a wind-pollinated turfgrass species, can efficiently disseminate herbicide resistance alleles via PMGF and movement of seeds and stolons. The genus Agrostis contains about 200 species, many of which are sexually compatible and produce naturally occurring hybrids and hybrids with species in the genus Polypogon. The self-incompatibility, extremely high outcrossing rate, and wind pollination in Italian ryegrass clearly point to PMGF as a major mechanism by which herbicide resistance alleles can spread across agricultural landscapes, resulting in abundant genetic variation within populations and low genetic differentiation among populations. Italian ryegrass can readily hybridize with perennial ryegrass and rigid ryegrass due to their similarity in chromosome numbers (2n = 14), resulting in interspecific gene exchange. Johnsongrass, barnyardgrass, and wild oats are self-pollinated species, so the potential for PMGF is relatively low and limited to short distances; however, seeds can easily shatter upon maturity before crop harvest, leading to wider dispersal. The occurrence of PMGF in reviewed grass weed species, even at a low rate, is greater than that of spontaneous mutations conferring herbicide resistance in weeds and thus can contribute to the spread of herbicide resistance alleles. This review indicates that the transfer of herbicide resistance alleles occurs under field conditions at varying levels depending on the grass weed species. Nomenclature: Barnyardgrass; Echinochloa crus-galli (L.) Beauv.; creeping bentgrass; Agrostis stolonifera L.; Italian ryegrass; Lolium perenne ssp. multiflorum (Lam.) Husnot; johnsongrass; Sorghum halepense (L.) Pers.; perennial ryegrass; Lolium perenne L.; rigid (annual) ryegrass; Lolium rigidum Geud.; sterile oat; Avena sterilis L.; wild oat; Avena fatua L.

Rigid ryegrass ( Lolium rigidum Gaud.) is one of the most troublesome weeds in Moroccan and Tunisian cereal crop fields. In total, 19 rigid ryegrass field populations were randomly selected in northern wheat crop areas of Morocco and Tunisia to examine the patterns of herbicide resistance to acetolactate synthase (ALS)- and acetyl-CoA carboxylase (ACCase)-inhibiting herbicides. Greenhouse experiments confirmed reduced sensitivity to ALS- and/or ACCase-inhibiting herbicides in all L. rigidum populations. The occurrence of target-site resistance (TSR) was tested using high-throughput genotyping. The advent of next-generation sequencing (NGS) has enabled easy identification of causal mutations and confirmed the presence of ALS and ACCase mutations at specific codons conferring TSR. Thirteen populations showed resistance to ALS-inhibiting herbicides associated with point mutations in positions Pro-197-Thr, Pro-197-Ser, Pro-197-Leu, Pro-197-Gln and Trp-574-Leu, while resistance to ACCase-inhibiting herbicides was detected in 18 populations in positions Asp-2078-Val, Trp-2027-Cys, Ile-1781-Leu, Gly-2096-Ala, and Ile-2041-Asn of the enzymes conferring TSR. Additionally, dose–response experiments with pyroxsulam applied after the inhibition of cytochrome P450 monooxygenase by malathion showed an increase in sensitivity in two out of seven highly resistant (HR) rigid ryegrass populations. This demonstrates the presence of non-target-site resistance (NTSR) in some ryegrass populations. Further evidence of NTSR was investigated in dose–response experiments with pyroxsulam, following pretreatment with the glutathione S -transferase (GST) inhibitor 4-chloro-7-nitrobenzoxadiazole (NBD-Cl), which partially reversed resistance in only a few individuals of two L. rigidum populations. Hence, our study confirms the existence of multiple and cross-resistance to ALS- and ACCase-inhibiting herbicides in L. rigidum from Morocco and Tunisia with both TSR and NTSR mechanisms. These results emphasize local resistance management as an important tool to detect and mitigate gene flow from rigid ryegrass populations where resistance has evolved.

Plants that release molecules affecting other plants are a source of potential bioherbicides. Silver wattle (Acacia dealbata Link), considered invasive worldwide, was found to be phytotoxic to various other plant species. Combining the search for alternative bioherbicides while reducing the spread of this invader by preventing seed formation is a good potential strategy to solve both agricultural and environmental problems. This study aimed to identify nonvolatile compounds from A. dealbata flowers and explore their phytotoxicity on the germination process and seedling and plant growth of lettuce (Lactuca sativa L.), wheat (Triticum aestivum L.), and rigid ryegrass (Lolium rigidum Gaudin). We identified methyl cinnamate and methyl anisate as potential phytotoxins in the extracts, but we used pure commercial molecules to conduct bioassays. Methyl cinnamate showed higher phytotoxicity than methyl anisate and was selected for further bioassays. Methyl cinnamate reduced guaiacol peroxidase activity by 57% and 85% in L. rigidum and lettuce, respectively, and α-amylase by 6% in L. rigidum. This compound also inhibited early stem and radicle growth of dicotyledonous lettuce (60% and 89%, respectively) and monocotyledonous L. rigidum (76% and 87%, respectively), both species having small seeds. However, wheat with a larger seed size was not affected by the phytotoxin. The results obtained indicate a potential bioherbicidal effect for methyl cinnamate, and its application might be useful in wheat crops infested by L. rigidum. We suggest that collecting A. dealbata flowers would prevent Acacia seed formation and thus play a role in invasive pest management, as well as serving as a source of potential herbicides to other species.

Crop allelopathy is a useful tool in integrated weed suppression. Although the abiotic stresses affect the produced secondary metabolites, the impact of soil salinity on crop allelopathic potential under field conditions has not been investigated. So, the effect of soil salinity on the allelopathic activity of 18 allelopathic barley (Hordeum vulgare L.) varieties was investigated. The allelopathy of barley grown on saline and non-saline soils was investigated during a 2-year field experiment using the Perlite-based bioassay with rigid ryegrass (Lolium rigidum L.) as plant indicator. The two main allelopathic substances in barley, gramine and hordenine, were determined using the high-performance liquid chromatography with tandem mass spectrometry. The results indicated that rigid ryegrass germination and root length were adversely affected by the barley aqueous extracts. The secondary metabolite gramine was detected in greater concentrations than those of hordenine. For most barley varieties, soil salinity reduced the phytotoxicity of aqueous extracts, as well as the concentration of gramine. However, gramine and hordenine were not highly correlated with the barley extract phytotoxicity, indicating the possible involvement of other allelopathic substances. In conclusion, the barley allelopathic potential probably decreases in soils with high salinity.

Chaff lining and chaff tramlining are harvest weed seed control (HWSC) systems that involve the concentration of chaff material containing weed seed into narrow (20 to 30 cm) rows between or on the harvester wheel tracks during harvest. These lines of chaff are left intact in the fields through subsequent cropping seasons in the assumption that the chaff environment is unfavorable for weed seed survival. The chaff row environment effect on weed seed survival was examined in field studies, and chaff response studies determined the influence of increasing amounts of chaff on weed seedling emergence. The objectives of these studies were to determine the influences of (1) chaff lines on the summer–autumn seed survival of selected weed species and (2) chaff type and amount on rigid ryegrass seedling emergence. There was frequently no difference (P > 0.05) in seed survival of four weed species (rigid ryegrass, wild oat, annual sowthistle, and turnip weed) when seeds were placed beneath or beside chaff lines. In one instance, wild oat seed survival was increased (P < 0.05) when seed were placed beneath compared to beside a chaff line. The pot studies determined that increasing amounts of chaff consistently resulted in decreasing numbers of rigid ryegrass seedlings emerging through chaff material. The suppression of emergence broadly followed a linear relationship in which there was approximately a 2.0% reduction in emergence with every 1,000 kg ha–1 increase in chaff material. This relationship was consistent across wheat, barley, canola, and lupin chaff types, indicating that the physical presence of the chaff was more important than chaff type. These studies suggested that chaff lines may not affect the survival over summer–autumn of the contained weed seeds but that the subsequent emergence of weed seedlings will be restricted by high amounts of chaff (>40,000 kg ha–1). Nomenclature: Annual sowthistle; Sonchus oleraceus L.; rigid ryegrass; Lolium rigidum Gaud.; turnip weed; Rapistrum rugosum (L.) All.; wild oat Avena fatua L.; barley; Hordeum vulgare L.; canola; Brassica napus L.; lupin; Lupinus angustifolius L.; wheat; Triticum aestivum L.

The Lolium genus encompasses many species that colonize a variety of disturbed and non-disturbed environments. Lolium perenne L. spp. perenne , L . perenne L. spp. multiflorum , and L . rigidum are of particular interest to weed scientists because of their ability to thrive in agricultural and non-agricultural areas. Herbicides are the main tool to control these weeds; however, Lolium spp. populations have evolved multiple- and cross-resistance to at least 14 herbicide mechanisms of action in more than 21 countries, with reports of multiple herbicide resistance to at least seven mechanisms of action in a single population. In this review, we summarize what is currently known about non-target-site resistance in Lolium spp. to acetyl CoA carboxylase, acetohydroxyacid synthase, microtubule assembly, photosystem II, 5-enolpyruvylshikimate-3-phosphate synthase, glutamine synthetase, very-long chain fatty acids, and photosystem I inhibitors. We suggest research topics that need to be addressed, as well as strategies to further our knowledge and uncover the mechanisms of non-target-site resistance in Lolium spp.

Multiple-herbicide resistance (MHR) in black-grass (Alopecurus myosuroides) and annual rye-grass (Lolium rigidum) is a global problem leading to a loss of chemical weed control in cereal crops. Although poorly understood, in common with multiple-drug resistance (MDR) in tumors, MHR is associated with an enhanced ability to detoxify xenobiotics. In humans, MDR is linked to the overexpression of a pi class glutathione transferase (GSTP1), which has both detoxification and signaling functions in promoting drug resistance. In both annual rye-grass and black-grass, MHR was also associated with the increased expression of an evolutionarily distinct plant phi (F) GSTF1 that had a restricted ability to detoxify herbicides. When the black-grass A. myosuroides (Am) Am GSTF1 was expressed in Arabidopsis thaliana, the transgenic plants acquired resistance to multiple herbicides and showed similar changes in their secondary, xenobiotic, and antioxidant metabolism to those determined in MHR weeds. Transcriptome array experiments showed that these changes in biochemistry were not due to changes in gene expression. Rather, Am GSTF1 exerted a direct regulatory control on metabolism that led to an accumulation of protective flavonoids. Further evidence for a key role for this protein in MHR was obtained by showing that the GSTP1- and MDR-inhibiting pharmacophore 4-chloro-7-nitro-benzoxadiazole was also active toward Am GSTF1 and helped restore herbicide control in MHR black-grass. These studies demonstrate a central role for specific GSTFs in MHR in weeds that has parallels with similar roles for unrelated GSTs in MDR in humans and shows their potential as targets for chemical intervention in resistant weed management.

The rate of herbicide resistance evolution in plants depends on fitness traits endowed by alleles in both the presence and absence (resistance cost) of herbicide selection. The effect of two Lolium rigidum spontaneous homozygous target-site resistance-endowing mutations (Ile-1781-Leu, Asp-2078-Gly) on both ACCase activity and various plant growth traits have been investigated here. Relative growth rate (RGR) and components (net assimilation rate, leaf area ratio), resource allocation to different organs, and growth responses in competition with a wheat crop were assessed. Unlike plants carrying the Ile-1781-Leu resistance mutation, plants homozygous for the Asp-2078-Gly mutation exhibited a significantly lower RGR (30%), which translated into lower allocation of biomass to roots, shoots, and leaves, and poor responses to plant competition. Both the negligible and significant growth reductions associated, respectively, with the Ile-1781-Leu and Asp-2078-Gly resistance mutations correlated with their impact on ACCase activity. Whereas the Ile-1781-Leu mutation showed no pleiotropic effects on ACCase kinetics, the Asp-2078-Gly mutation led to a significant reduction in ACCase activity. The impaired growth traits are discussed in the context of resistance costs and the effects of each resistance allele on ACCase activity. Similar effects of these two particular ACCase mutations on the ACCase activity of Alopecurus myosuroides were also confirmed.

Dormancy in Lolium rigidum (annual ryegrass) seeds can be alleviated by warm stratification in the dark or by application of fluridone, an inhibitor of plant abscisic acid (ABA) biosynthesis via phytoene desaturase. However, germination and absolute ABA concentration are not particularly strongly correlated. The aim of this study was to determine if cytokinins of both plant and bacterial origin are involved in mediating dormancy status and in the response to fluridone. Seeds with normal or greatly decreased (by dry heat pre-treatment) bacterial populations were stratified in the light or dark and in the presence or absence of fluridone in order to modify their dormancy status. Germination was assessed and seed cytokinin concentration and composition were measured in embryo-containing or embryo-free seed portions. Seeds lacking bacteria were no longer able to lose dormancy in the dark unless supplied with exogenous gibberellin or fluridone. Although these seeds showed a dramatic switch from active cytokinin free bases to O-glucosylated storage forms, the concentrations of individual cytokinin species were only weakly correlated to dormancy status. However, cytokinins of apparently bacterial origin were affected by fluridone and light treatment of the seeds. It is probable that resident microflora contribute to dormancy status in L. rigidum seeds via a complex interaction between hormones of both plant and bacterial origin. This interaction needs to be taken into account in studies on endogenous seed hormones or the response of seeds to plant growth regulators.

The legacy of deliberate and accidental introductions of invasive alien species to Australia has had a hefty economic toll, yet quantifying the magnitude of the costs associated with direct loss and damage, as well as for management interventions, remains elusive. This is because the reliability of cost estimates and under-sampling have not been determined. We provide the first detailed analysis of the reported costs associated with invasive species to the Australian economy since the 1960s, based on the recently published InvaCost database and supplementary information, for a total of 2078 unique cost entries. Since the 1960s, Australia has spent or incurred losses totalling at least US $298.58 billion (2017 value) or AU$ 389.59 billion (2017 average exchange rate) from invasive species. However, this is an underestimate given that costs rise as the number of estimates increases following a power law. There was an average 1.8–6.3-fold increase in the total costs per decade since the 1970s to the present, producing estimated costs of US $6.09–57.91 billion year -1 (all costs combined) or US$ 225.31 million–6.84 billion year -1 (observed, highly reliable costs only). Costs arising from plant species were the highest among kingdoms (US $151.68 billion), although most of the costs were not attributable to single species. Of the identified weedy species, the costliest were annual ryegrass ( Lolium rigidum ), parthenium ( Parthenium hysterophorus ) and ragwort ( Senecio jacobaea ). The four costliest classes were mammals (US$ 48.63 billion), insects (US $11.95 billion), eudicots (US$ 4.10 billion) and monocots (US$1.92 billion). The three costliest species were all animals – cats ( Felis catus ), rabbits ( Oryctolagus cuniculus ) and red imported fire ants ( Solenopsis invicta ). Each State/Territory had a different suite of major costs by species, but with most (3–62%) costs derived from one to three species per political unit. Most (61%) of the reported costs applied to multiple environments and 73% of the total pertained to direct damage or loss compared to management costs only, with both of these findings reflecting the availability of data. Rising incursions of invasive species will continue to have substantial costs for the Australian economy, but with better investment, standardised assessments and reporting and coordinated interventions (including eradications), some of these costs could be substantially reduced.