Management of kudzu by the bioherbicide,Myrothecium verrucaria,herbicides and integrated control programmes

Replicated field plots were established and monitored for two years to evaluate management practices for kudzu. The bioherbicidal plant pathogen, Myrothecium verrucaria, several herbicides and a variety of integrated control programmes achieved a high level of kudzu suppression, although no system tested reliably achieved eradication in this time frame.     

Bioherbicidal activity from washed spores of <Emphasis Type="Italic">Myrothecium verrucaria</Emphasis>

The fungal plant pathogen, Myrothecium verrucaria, is highly virulent to several important weed species and has potential utility as a bioherbicide. However the production of macrocyclic trichothecene mycotoxins by this fungus presents significant safety concerns. It was discovered that trichothecenes are removed from M. verrucaria spores by repeated washes with water. These washed spores retained bioherbicidal efficacy against kudzu when tested in field trials and on sicklepod when tested under greenhouse conditions. Changes in the growth medium combined with washing spores with water resulted in greater than 95% reduction in roridin A and verrucarin A. Washing spores reduced trichothecene concentrations in spore preparations with no significant effect on plant biomass reduction, thus demonstrating the possibility of M. verrucaria formulations with improved safety to researchers, producers and applicators.     

Effect of turfgrass mowing height on biocontrol of dandelion with Sclerotinia minor

The fungus, Sclerotinia minor Jagger is under development as a bioherbicide for control of dandelion and many broadleaf weeds in turfgrass environments. The effect of S. minor on dandelion survival was evaluated under different mowing heights and compared with a commonly used herbicide Killex™. In the greenhouse, the onset of symptoms was more rapid, foliar damage was more severe, and the reduction of aboveground biomass and root biomass was greater for the bioherbicide than the herbicide. The bioherbicide reduced root biomass ≥10-fold compared with untreated plants. Under high weed infestation levels in the field, S. minor caused a greater initial reduction of dandelion density than did the herbicide during the 2-week post-application period, although reductions were greater in herbicide treated plots by 6 weeks after application. Over the growing season, S. minor and the herbicide had similar suppressive effects on dandelion density except under the closest mowing height (3-5 cm). After treatment, close mowing favored dandelion seedling recruitment and the biocontrol had no residual activity. Survival of dandelion roots was significantly less after spring than fall treatment of S. minor and season long mowing at the close height significantly reduce root survival. Close mowing may be detrimental for S. minor applications on heavily infested domestic lawns and amenity grassland areas.     

Submerged culture of a mycelial formulation of a bioherbicidal strain of <Emphasis Type="Italic">Myrothecium </Emphasis><Emphasis Type="Italic">verrucaria</Emphasis> with mitigated mycotoxin production

A mycelial formulation of the fungus Myrothecium verrucaria (IMI 361690) containing 0.20% Silwet L-77 surfactant was found to be highly efficacious in controlling the exotic invasive weed kudzu. The mycelium can be rapidly (48–72 h) produced in several media, including an inexpensive soy flour–corn meal medium. Mycelial yields were 2, 10, and 25 g dry weight l⁻¹ in Czapek-Dox, Richard’s V-8, and soy flour–corn meal media, respectively. Scale-up production in soy flour–corn meal medium using laboratory fermenters (10–25 l), resulted in a mycelial formulation that caused 90% mortality of naturally-occurring mature (0.9–1.0 m in height) kudzu within 48 h after application in field experiments. HPLC analyses revealed that the mycelium produced in this liquid culture contained no detectable amounts of the trichothecene mycotoxins roridin A and verrucarin A (limit of detection 2 μg ml⁻¹). This has resulted in a safer, yet effective bioherbicidal product. We anticipate that these findings should improve the probability of EPA registration and subsequent commercial development of this bioherbicide.     

Effectiveness and technological feasibility of bioherbicide candidates for biocontrol of Green Foxtail (Setaria viridis)

Green foxtail (Setaria viridis), one of the most common and troublesome weeds worldwide, is becoming very difficult to manage because of the lack of registered herbicides and the appearance of herbicide-resistant populations. Among the new and possible environment-friendly strategies, the use of biological control methods seems to have potential. Drechslera gigantea, Exserohilum rostratum and Exserohilum longirostratum have previously proved to be promising bioherbicide agents against several grass weeds in field trials. While previous studies have established the susceptibility of S. viridis under greenhouse conditions, so far no attempt has been made to establish the effectiveness and feasibility of these fungi as bioherbicides for green foxtail. When spore suspensions were applied as foliar sprays to green foxtail seedlings in a greenhouse, all three fungi caused severe damage by 1 day after application, and seedlings in most cases died within 1 week. The fungi were compatible with several agro-chemicals and host specific when tested against major vegetable crop species grown in the Mediterranean. The demonstrated technological feasibility of producing large amounts of quickly germinable conidia (i.e., asexual spores) on inexpensive solid media increases the potential of these fungi to be used as bioherbicides.     

Myrothecium verrucaria isolates and formulations as bioherbicide agents for kudzu

The fungus Myrothecium verrucaria (MV) has previously been shown to have potential as a bioherbicide for kudzu (Pueraria lobata) control. It has also been shown that MV wild-type (MV-wt) often forms sectors, when grown on various nutrient media. Experiments compared MV-wt and MV sector efficacy when grown on agar or on rice grains. In greenhouse evaluations of sectors, applied as foliar sprays in water or in other formulations (corn oil, surfactant, and corn oil plus surfactant) for efficacy against kudzu seedlings, some sectors possessed bioherbicidal activity equal that of MV-wt, but others exhibited lower activity. Without a dew period, aqueous formulations of MV-wt, a yellow sector, and a white sector provided zero control, but all three isolates were active without a dew period when formulated in corn oil, Silwet L-77 surfactant, and in surfactant plus corn oil. Generally, the yellow sector was less effective than the other two isolates in any formulations, and the MV-wt and white sector provided approximately 100% mortality of the test plants. Dew (10 h) increased weed control to 100, 33, and 65%, respectively, for MV-wt, the yellow sector and the white sector. All isolates provided nearly 100% control in the oil and surfactant formulations with a dew period compared to treatments receiving no dew. Soil incorporation studies were also performed to compare MV-wt efficacy of preparations grown on agar versus growth on rice grains. Higher efficacies (1.75-3.3-fold increase) were obtained from rice grain preparations compared to preparations grown on agar, when preparations were incorporated at several rates into soil prior to planting. Cell-free extracts of the MV-rice cultures were also phytotoxic to kudzu seedlings up to the eight- to 10-leaf growth stage. Thus, formulation, growth media, and the application method are important determinants in the efficacy of MV and MV sectors on kudzu seedlings.     

Weed development in spring wheat after contrasting soil tillage and nitrogen management

Soil tillage and nitrogen (N) management effects on weed species composition were evaluated in 2013 and 2014 on a clayey soil after 5‐years of organic management at the Royal Agricultural University's Harnhill Manor Farm, UK. Three tillage systems – Conventional Tillage (CT), and High and Low Intensity Non‐inversion Tillage (HINiT & LINiT) – were compared at four N fertiliser rates of 0, 70, 140 and 210 kg N ha^?1. Broad‐spectrum herbicide was applied before soil operations across the site in both years. Previous organic management legacy of high weed biomass promoted greater weed prevalence in 2013 while 2‐years of herbicide inclusion reduced weed biomass. Contrasting weather conditions across the seasons affected weed incidence. In the 2014 wet season, early weed dry weight (DM) was higher under HINiT than CT and LINiT, while no differences were observed in the 2013 dry year. At midseason, weed DM was higher under HINiT than CT and LINiT in both years, which was related to higher DM of the dominant weeds Stellaria media (L.) Vill. and Sinapis arvensis L. Grass weed DM was higher under non‐inversion tillage than CT. N fertilisation increased midseason total weed DM and weed prevalence at harvest. Spring wheat yield was the highest under CT while LINiT produced 17% higher yields than HINiT. Despite higher but still tolerable weed prevalence under both non‐inversion tillage systems and with the application of N, weeds alone was not the only yield‐limiting factor. However, results show that CT is the most reliable option for weed control in changing weather, while N fertilisation rates needs to be considered.     

Towards the biocontrol of bindweeds with a mycoherbicide

Within the framework of the European COST Action 816, afive-year collaboration between scientists from five Europeancountries has made an important contribution to biologicalcontrol of field and hedge bindweeds (Convolvulus arvensis andCalystegia sepium, respectively). A fungus Stagonosporaconvolvuli strain LA39, able to infect both field and hedgebindweed, was found in the UK and its biocontrol efficacyimproved by optimising mass production, formulation and storagetechniques. This fungus controlled bindweeds in both a cemeteryand in maize crops. Its use fits best in an integrated pestmanagement system where a green cover controls most of the weedsexcept the bindweeds. DNA marker analyses indicate that thefungus reproduces sexually, which could be used to furtherimprove this mycoherbicide. In addition, the insect Melanagromyzaalbocilia, which itself exhibits biocontrol potential againstbindweeds, may be used in combination with LA39 to improve theability of the fungus to penetrate the stem of bindweeds.Overall, the results suggest that S. convolvuli LA39 haspromising potential as a bioherbicide for control of field andhedge bindweed.     

Development of strain specific molecular markers for the Sclerotinia minor bioherbicide strain IMI 344141

The plant pathogenic fungus, Sclerotinia minor IMI 344141, has been developed as a bioherbicide for broadleaf weed control in turfgrass and a means to differentiate this biocontrol agent from like organisms is required. A strain specific molecular marker was developed to detect and monitor the Sclerotinia minor IMI 344141 bioherbicide strain. The method was based on polymerase chain reaction (PCR) amplification of two sequence-characterized amplified regions (SCAR) primer pairs for a first round PCR, and another two sets of nested primers was used for a second round PCR if higher sensitivity was needed. Sclerotinia minor IMI 344141 was successfully traced from both pure cultures and environmental samples originating from bioherbicide-released field trials. DNA of the S. minor bioherbicide isolate IMI 344141 was detected in the soil 2 months after application, but was not detected in the 3- and 9-month samples after application. When applied as a bioherbicide, S. minor (IMI 344141) did not persist into the following spring season in turf environments. This molecular detection method provides a mechanism to distinguish this isolate from related organisms and a tool to monitor behavior of the biocontrol agent S. minor IMI 344141 in nature, particularly in soil.     

Preservation, mass production and storage ofStagonospora convolvuli, a bioherbicidecandidate for field bindweed (Convolvulusarvensis)

Different solid substrates were investigated as spore production methods for Stagonospora convolvulistrain LA39, a potential bioherbicide for field bindweed (Convolvulusarvensis L.). Up to 4 × 10⁸ spores/g of substratewere yielded on cous-cous (cracked hard wheat). Thespores were as pathogenic as those grown on artificial medium (V-8-juice agar). The air-drying on kaolin and storage at 3 °C kept spores viable and pathogenic for 180 days. Spore germination exceeded70% for the first 140 days and then declined to 50%after 175 days. Less than 5% of spores were still viable after 17 months. The preservation of stock cultures in 10% glycerine at −80 ° C and in liquid nitrogen did not affect viability orpathogenicity of the spores. This revised version was published online in July 2006 with corrections to the Cover Date.     

Prospects for the Management of Invasive Alien Weeds Using Co-Evolved Fungal Pathogens: A Latin American Perspective

Invasive alien weeds pose a serious threat to the biodiversity of natural ecosystems and a significant constraint to agricultural production worldwide. The use of co-evolved natural enemies, a strategy referred to as classical biological control (CBC), has proven to be a potentially efficacious, cost-effective, and safe option for the management of alien weeds. An analysis of CBC of invasive weeds in Latin America is presented, which shows that only 5% of the worldwide releases of agents, overwhelmingly arthropod, have been in this region. Fungal pathogens are increasingly being considered in CBC programmes, and there are now 11 examples of Latin American fungi having been released as biocontrol agents in other regions of the world. In contrast, only three weed pathogens have been deliberately released in the region. Possible reasons for the paucity of CBC programmes in Latin America are presented, despite the presence of a significant number of alien weed species (60 are listed). An analysis of these weeds reveals that many of them could be amenable to control using natural enemies, including nine weed species for which CBC programmes have been successfully implemented elsewhere in the world. In addition, for many of these 60 species, a co-evolved and damaging mycobiota has already been recorded. The prospects for management of invasive alien weeds in Latin America, using co-evolved fungal pathogens, are assessed with particular reference to selected species from the genera Ambrosia, Broussonetia, Calotropis, Commelina, Cyperus, Dichrostachys, Echinochloa, Pittosporum, Rottboellia, Rubus, Sonchus and Ulex. This revised version was published online in July 2006 with corrections to the Cover Date.     

Microencapsulated Spodoptera frugiperda nucleopolyhedrovirus: insecticidal activity and effect on arthropod populations in maize

Excessive use of chemical insecticides to control Spodoptera frugiperda in maize (Zea mays L.) crops in Colombia has resulted in pest resistance and environmental contamination. A Colombian multicapsid nucleopolyhedrovirus (SfMNPV) was produced in laboratory reared larvae and formulated as a wettable powder by microencapsulating the viral occlusion bodies (OBs) with a methacrylic acid polymer. Microencapsulated OBs were tested for insecticidal activity in laboratory, greenhouse and field bioassays, and were then sprayed onto a commercial crop where their effect on arthropod populations was estimated. In laboratory bioassay, microencapsulated and non-microencapsulated OBs caused a larval mortality of 98 and 96%, respectively. The percent of plants showing recent feeding damage in a greenhouse experiment oscillated between 2.5 and 7.5% when OBs were applied compared to 70% observed in the control. In two field trials, the highest dose of 1.5×10¹² OBs/ha (1500 g/ha) was selected as the recommended rate for field application in maize crops, with efficacies higher than 70%. Total arthropod populations were negatively affected by chemical pesticides; whereas the OBs formulation did not affect the diversity of these organisms. Our results indicate that microencapsulated SfMNPV has the potential to be commercialised as an alternative to substitute or complement chemical insecticides in fall armyworm integrated management programmes.     

Acetone-butanol-ethanol fermentation and isoflavone extraction using kudzu roots

The economics of Acetone-butanol-ethanol (ABE) fermentation is greatly affected by raw materials, and the use of readily available starchy materials from marginal farming lands could be a viable option for reducing costs. Kudzu, a rapidly growing perennial leguminous vine, has been planted on marginal farming land and widely distributed in Asia and America. This study investigated ABE fermentation by C. acetobutylicum ATCC 824 using kudzu roots and isoflavone extraction from kudzu fermentation residue (KFR). The kudzu roots could be used as a sole substrate for ABE fermentation without nutritional supplements. Batch culture containing 140 g kudzu/L produced 17.99 ± 1.08 g/L solvent (ABE), including 11.20 ± 0.79 g/L butanol, 5.54 ± 0.20 g/L acetone, and 1.15 ± 0.09 g/L ethanol, with a productivity of 0.19 g/(L/h) and a yield of 0.33 g solvent/g sugar after 96 h of fermentation. Isoflavone yield extracted from KFR was 1.90/100 g KFR, approximately 48% higher compared with that extracted from raw kudzu. A kinetic analysis of the extraction process showed that both the isoflavone yield and the extraction rate obtained from KFR were higher than the corresponding values obtained from raw kudzu. These results indicate that kudzu may provide a new potential raw material for ABE production and the process of ABE fermentation integrated with isoflavone extraction may provide a new way to reduce fermentable substrate costs.     

Response of Acroptilon repens to simulated herbivory and soil disturbance

Acroptilon repens is an invasive weed in North America but also causes problems in disturbed habitats in its native range in Asia. In order to test the effect of simulated biological control and soil disturbance on established A. repens patches and the competing vegetation, two levels of shoot clipping as well as soil tillage were imposed on A. repens patches in an undisturbed meadow and at two fallowland sites in the native range of the weed. At the meadow site, 2 years of partial clipping of shoots and of soil tillage had no influence on A. repens performance, while soil tillage significantly reduced the above‐ground biomass of the competing vegetation. At the fallowland sites, which had been continuously cultivated for several years prior to the experiment, A. repens shoot density, biomass and number of seed heads were significantly higher in the undisturbed control than in the tillage plots. The total number of seed heads per unit area increased with shoot density up to 200 shoots m^?2. These results indicate that A. repens has considerable regrowth capacities that allow established patches to tolerate substantial losses of above‐ground biomass and that the competitive ability of A. repens is favoured both when soil disturbance is imposed on previously undisturbed sites, as well as when repeated soil disturbance is abandoned. The only promising nonchemical herbicide‐based approach to reduce the competitive ability and seed output of A. repens appears to be a long‐term management that enhances the interspecific plant competition by reducing soil disturbance and selectively damaging A. repens.     

Enzyme Responses of <Emphasis Type="Italic">Abutilon Theophrasti</Emphasis> in an Enhanced Biocontrol System

Weak virulence has limited the development of Colletorichum coccodes (Wallr.) Hughes (DAOM 183088) as a bioherbicide for Abutilon theophrasti Medik. (velvetleaf) control. This study examines the role of chemical synergy to suppress host defense mechanisms. Dry weight of A. theophrasti was reduced by treatment with C. coccodes in a tank-mix with 0.25 kg a.i. ha⁻¹ bentazon more than by the treatment with C. coccodes or bentazon alone, while the effect of a sub-lethal rate of glyphosate was not significant. C. coccodes did not affect phenylalanine ammonia lyase (PAL) activity in infected leaves of A. theophrasti, whereas PAL activity was significantly inhibited by the presence of bentazon 5 days after treatment. The inhibition was stronger when bentazon was applied alone than when bentazon was applied with C. coccodes. Treatment with glyphosate or a mixture of glyphosate and C. coccodes did not affect PAL activity. Peroxidase activity was strongly induced by C. coccodes treatment and increased over time. Peroxidase activity was not induced by 0.25 kg a.i. ha⁻¹ bentazon alone. However, when bentazon was applied in combination with C. coccodes, it prevented the activation of peroxidase by C. coccodes infection. Treatment with 0.2 kg a.i. ha⁻¹ glyphosate did not affect peroxidase activity. These results suggest that these enzymes are involved in the resistance mechanism of A. theophrasti, and the synergistic effect of bentazon on C. coccodes efficacy as a bioherbicide may result from suppressing the activation of these defense-related enzymes.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Effects of weed harrowing frequency on beneficial arthropods,plants and crop yield

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A genomic approach to isoflavone biosynthesis in kudzu (<Emphasis Type="Italic">Pueraria lobata</Emphasis>)

Roots of kudzu (Pueraria lobata) are a rich source of isoflavone O- and C-glycosides. Although O-glycosylation of (iso)flavonoids has been well characterized at the molecular level, no plant isoflavonoid C-glycosyltransferase genes have yet been isolated. To address the biosynthesis of kudzu isoflavonoids, we generated 6,365 high-quality expressed sequence tags (ESTs) from a subtraction cDNA library constructed using RNA from roots that differentially accumulate puerarin. The ESTs were clustered into 722 TCs and 3,913 singletons, from which 15 family I glycosyltransferases (UGTs) were identified. Hierarchical clustering analysis of the expression patterns of these UGTs with isoflavone synthase (IFS) in a range of tissues identified UGTs with potential functions in isoflavone glycosylation. The open reading frames of these UGTs were expressed in E. coli for functional analysis, and one was shown to preferentially glycosylate isoflavones at the 7-O-position. In addition, ESTs corresponding to chalcone synthase, chalcone reductase, chalcone isomerase (CHI) and 2-hydroxyisoflavanone dehydratase were identified. Recombinant CHI proteins had high activities with both 6′-deoxy- and 6′-hydroxy chalcones, typical of Type II CHIs. Establishment of this EST database and identification of genes associated with kudzu isoflavone biosynthesis and glycosylation provide a new resource for metabolic engineering of bioactive kudzu isoflavones.     

Population statistics and biological traits of Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae) affected by LC30 of thiamethoxam and pirimicarb

Although selective pesticides are recommended to use in integrated pest management (IPM) programmes, they could adversely affect biological agents. So, the aim of the current study was to investigate the LC₃₀ of pirimicarb and thiamethoxam on Hippodamia variegata. Two pesticides, pirimicarb and thiamethoxam, when applied at concentrations of 1522.8 and 251.3?mg (ai)/L, respectively, on the third instar larvae of the insect, produced 30% mortality. Also, results showed that neither pirimicarb nor thiamethoxam affect stage age distribution (c_x), fecundity (eggs/female) and adult development time. These two pesticides extended preadult duration significantly (p?<?0.0001, F?=?31.22, df?=?122). Also, survival rate (l_x) and age-specific reproductive value (v_x) decreased and some changes in age-specific life expectancy (e_x) happened. Generally, more adverse effects were found in the population treated by thiamethoxam. The results showed that pirimicarb and thiamethoxam have potential to harm the predatory ladybird in IPM programme, though in sublethal doses.     

Biological Control of Kudzu ( Pueraria lobata ) with an Isolate of Myrothecium verrucaria

An isolate of the fungus Myrothecium verrucaria ( MV ) was evaluated for biocontrol potential against kudzu ( Pueraria lobata ). In greenhouse tests, MV was highly virulent against kudzu in the absence of dew when conidia were formulated in 0.2% Silwet L-77 surfactant (SW). Inoculum concentrations ≥2 ×10 7 conidia ml -1 were required to satisfactorily control plants in the third leaf stage and larger. In controlled environment experiments, kudzu mortality was greater at higher temperatures (25-40°C) than at lower temperatures (10-20°C), although pathogenesis and mortality occurred at all temperatures tested. In field tests, transplanted kudzu seedlings in the 2-3 leaf growth stage treated with MV at 2 ×10 7 conidia ml -1 in 0.2% SW, exhibited leaf and stem necrosis within 24 h following inoculation, with mortality occurring within 96 h. After 7 days, 100% of inoculated kudzu plants were killed in plots treated with the fungus/surfactant mixtures. Similar results were observed in a naturally occurring kudzu population, where 100% control occurred within 14 days after inoculation with 2 ×10 7 conidia ml -1 in 0.2% SW. In summary, MV effectively controlled kudzu in the absence of dew over a wide range of physical and environmental conditions and under field conditions. These results indicate that, when properly formulated, MV has potential as a valuable bioherbicide for controlling kudzu.