Can synthetic pesticides be replaced with biologically-based alternatives?—an industry perspective

Agricultural chemical companies have invested in the discovery and development of biological pesticides to complement synthetic pesticides for the control of insects, diseases, and weeds on agronomic and horticultural crops. For plant disease control, companies envisage biological fungicides entering markets where they have the best chance of performing and which are most receptive to using biological control methods. Fewer regulatory requirements can mean faster registration for a biological than a synthetic pesticide. However, industry’s requirements for competitive performance, effective formulations, and economic production can mean significant investments in time and money for a biological pesticide, although total investment may be less than for a synthetic pesticide. One biocontrol project in which industry has invested is baculoviruses for insect control. Insect baculoviruses, genetically modified to kill insects faster than wild-type viruses, are attractive biocontrol agents because their selectivity to insect pests and safety to beneficial insects and mammals enable them to compete with synthetic insecticides. Industry is looking for similar biocontrol opportunities in disease control. Biocontrol agents for seedling disease, root rot, and postharvest disease control have been registered by the EPA and are trying to compete with synthetic fungicides for market share. To date, effective biocontrol agents have not been identified for the control of serious foliar diseases, such as grape downy mildew, potato late blight, wheat powdery mildew, and apple scab. Farmers must rely on synthetic fungicides and agronomic methods to control these diseases for the foreseeable future. Received 06 February 1997/ Accepted in revised form 01 June 1997     

The effects of flooding,plant traits,and predation on purple loosestrife leaf‐beetles

Classical biological weed control is based on the premise that introducing specialized natural enemies from the native range re‐establishes herbivore control of plant invaders, ultimately leading to negative population growth rates. Evidence from past biocontrol programs suggests that herbivores are not solely responsible for shaping plant demography. Diverse environmental conditions in the introduced range may not only affect demography, but also influence top‐down control of target plants. We investigated how flooding affects impacts of predators (top‐down) and plant quality (bottom‐up) on performance of two leaf‐beetles, Galerucella calmariensis L. and Galerucella pusilla Duftschmid (Coleoptera: Chrysomelidae: Galerucini), released in North America as biocontrol agents of purple loosestrife, Lythrum salicaria L. (Lythraceae). Predation and flooding regime have been linked to low leaf‐beetle recruitment at sites where insects failed to attain outbreak populations. Predator exclusion experiments at adjacent flooded and non‐flooded sites indicated a positive effect of flooding on leaf‐beetle survival for all developmental stages, whereas predator exposure had little effect. There was no difference in predation rates at sites with successful or failed purple loosestrife control, questioning the importance of predation in limiting growth and impact of these biocontrol agents’ populations. Effect of flooding on purple loosestrife quality was evaluated in a common garden study where plants were grown under different flooding treatments. Plants grown in flooded soil had higher water content and lower tannic acid concentration than plants grown in well‐drained soil. Consistent with field observations, leaf‐beetle oviposition rate and survival were higher on flooded plants. Results indicate that both bottom‐up and top‐down forces operate on Galerucella populations, yet their relative strength is mediated by flooding regime. Ignoring intricacies of plant‐herbivore and trophic interactions in the introduced range appears to be a major handicap for the improvement of weed biocontrol programs.     

Review of entomopathogenic fungi and nematodes as biological control agents of tephritid fruit flies: current status and a future vision

Several fruit fly species (Diptera: Tephritidae) are invasive pests that damage the quality of fruits in horticultural crops and cause significant value losses worldwide. Management of fruit flies mainly depends on conventional insecticides. Unfortunately, the application of synthetic insecticides has caused environmental pollution, risks for humans and animals, and development of resistance. Furthermore, controlling fruit flies by applying synthetic insecticides is challenging because fruit containing third instars often fall from the tree – subsequently the larvae leave the decaying fruits and pupate in the soil. Consequently, both larvae and pupae are protected from surface-applied insecticides in fruits and soil. So, there is a pressing need for more eco-friendly and selective control measures with new modes of action. Among such measures are entomopathogenic fungi (EPFs) and nematodes (EPNs). I gathered knowledge on past and present research about EPFs and EPNs as biocontrol agents against fruit flies to investigate approaches that may improve their capacities. I also highlighted several recommendations that may help future field studies on the suppression of fruit fly populations by EPFs and EPNs.     

Conundrums of a complex vector for invasive species control: a detailed examination of the horticultural industry

Historically the horticultural industry has transformed the US landscape through intentional cultivar introductions and unintentional introductions of weeds, insects and plant diseases. While it has been demonstrated that the horticultural industry, in particular the ornamental subsector, is an important vector for the introduction and dispersal of invasive species, known invasive plants continue to be sold while new cultivars are introduced at an ever increasing rate. This study examines the horticultural trade as a vector for invasive species, its agents, and characterizes the complexity of the distribution channel. Numerous factors have contributed to the recent expansion in marketed cultivars, including technological, industry growth, and marketing developments. The result has been an increased and sophisticated consumer demand with a corresponding aggressive scouring of the planet for new crops, many of which are introduced into the market without sufficient testing for invasive tendencies. Traditional approaches to invasive horticultural crop control (regulation, self-regulation), which target players in the distribution channel before and/or after cultivar release, have had limited effectiveness and buy-in because these approaches do not address the industry’s complexities and economic incentives. Involvement and education of consumers may provide better oversight outcomes by addressing the moral hazard problem while acknowledging the key characteristics of the industry.     

Significance of ecology in the development of biocontrol agents against soil‐borne plant pathogens

Approaches to the development of inoculant biocontrol agents (BCAs) of soil‐borne pathogens are discussed. Based on an analysis of the success of Peniophora (Phlebia) gigantea and Agrobacterium radiobacter it is argued that most subsequent attempts to develop inoculant BCAs have failed because the organisms were selected for in vitro antagonism but were ecologically unsuited to the environments where pathogens grow. The reported modes of action of BCAs are reviewed and in some cases reinterpreted. It is argued that antibiosis and some types of mycoparasitism have not been shown to be direct mechanisms of biocontrol in vivo; they might, instead, facilitate competition for substrates or sites as the primary mechanism of control. The ecology of BCAs in general is reviewed, with emphasis on interactions in microsites and the design of appropriate screening strategies. Specific examples are used to illustrate these points.     

Biological control of microbial infections and cancer in humans: Historical use to future potential

Prior to the advent of antibiotics, live organisms were used directly in attempts to control microbial infections and cure cancers. Examples of such biological control included bacteriotherapy, bacteriophage therapy, malaria therapy, probiotics and the use of living maggots. In all cases, the organisms themselves, rather than products of their metabolism, were used as the potentially curative agents. The history of the use of biocontrol agents in the treatment of human infections and cancer is discussed here in relation to more recent examples of the use of this approach. Modern studies suggest that the use of biological control in the treatment of human infections may be worth re‐evaluating in the light of the increasing world‐wide occurrence of antibiotic‐resistant bacteria and the opportunities provided by recent developments in gene technology.     

A new formulation system for the application of biocontrol fungi to soil

A new biocontrol formulation system was devised that does not require sterile conditions during preparation. It involves mixing vermiculite and powdered wheat bran with wet or dry fermentor biomass of Trichoderma spp. or Gliocladium virens, moistening with 0.05 N HCl, and drying the mixture. Before application to soil, the preparation (VBA‐FB) is activated by re‐moistening with 0.05 N HCl and incubated at room temperature for 2–3 days to stimulate development of young hyphae of the biocontrol fungus. Populations of biocontrol fungi proliferated to greater than 10⁷ colony‐forming units (cfu) per g of soil when activated VBA‐FB was added to soil. In soil artificially infested with Rhizoctonia solani, seven isolates of the 14 studied added as VBA‐FB reduced survival and 12 reduced saprophytic growth of the pathogen. Of these, two isolates of T. hamatum (TRI‐4, Tm‐23) and one of T. harzianum (Th‐87) were the most effective. Preparations formulated with either wet or dry biomass effectively reduced pathogen survival, but activated VBA‐FB was more effective than non‐activated VBA‐FB. Storage of VBA‐FB at 25°C for 24 weeks before activation reduced viability of isolates considerably more than storage at 5°C for 24 weeks. In addition, VBA‐FB stored at 5°C before activation more effectively reduced survival of R. solani than VBA‐FB stored at 25° C. Survival of R. solani was reduced by activated VBA‐FB applied to several soil types (sandy loam, sandy clay loam, clay). Some nitrogen fertilizers increased the efficacy of VBA‐FB preparations of several isolates.     

Soil solarisation, amendments and bio-control agents for the control of Macrophomina phaseolina and Fusarium oxysporum f.sp. cumini in aridisols

The effects of soil solarisation, residue incorporation, summer irrigation and biocontrol agents singly or in combination on survival of Macrophomina phaseolina and Fusarium oxysporum f.sp. cumini were ascertained in the 2000 and 2001 summer seasons. In amended plots, temperature increased by 2.5°C over non‐amended plots (42–51°C) at various soil depths. Combining amendments and soil solarisation elevated the soil temperatures by 0.5–5°C and 2.5–13.0°C compared to non‐amended solarised and non‐solarised plots, respectively. These treatment combinations significantly reduced M. phaseolina and Fusarium propagules compared to control. Of these, combining mustard pod residues with soil solarisation almost eliminated viable propagules of both the pathogens at 0–30 cm soil depth. However, a combination of mustard pod residue and oil‐cake (2.5 + 0.5 ton ha^?1) with only one summer irrigation also caused pronounced reduction in pathogenic propagules, which was equal to that recorded in non‐amended solarised plots. The effect of surviving propagules of M. phaseolina and Fusarium on incidence of dry root rot on clusterbean and wilt on cumin was studied in subsequent rainy and winter seasons, respectively. Significant reductions in both diseases were recorded in residue and biocontrol amended plots with or without polyethylene mulching compared to non‐amended control. The lowest plant mortality in both the crops was recorded in mustard residue amended solarised plots in a two year field experiment. However, the disease indices in the plots having a combination of mustard residues and oil‐cake amendment with one summer irrigation was equal to that achieved in the treatment having polyethylene mulching. These results suggest that in hot arid regions use of Brassica residues can be a practical and feasible substitute for polyethylene mulching in managing soil‐borne diseases.     

Review: predatory soil mites as biocontrol agents of above- and below-ground plant pests

Biological pest control is becoming increasingly important for sustainable agriculture. Although many species of natural enemies are already being used commercially, efficient biological control of various pests is still lacking, and there is a need for more biocontrol agents. In this review, we focus on predatory soil mites, their role as natural enemies, and their biocontrol potential, mainly in vegetable and ornamental crops, with an emphasis on greenhouse systems. These predators are still underrepresented in biological control, but have several advantages compared to predators living on above-ground plant parts. For example, predatory soil mites are often easy and affordable to mass rear, as most of them are generalist predators, which also means that they may be used against various pests and can survive periods of pest scarcity by feeding on alternative prey or food. Many of them can also endure unfavourable conditions, making it easier for them to establish in various crops. Based on the current literature, we show that they have potential to control a variety of pests, both in greenhouses and in the field. However, more research is needed to fully understand and appreciate their potential as biocontrol agents. We review and discuss several methods to increase their efficiency, such as supplying them with alternative food and changing soil/litter structure to enable persistence of their populations. We conclude that predatory soil mites deserve more attention in future studies to increase their application in agricultural crops.

     

Molecular and biochemical aspects of rhizobacterial ecology with emphasis on biological control

The rhizosphere is the narrow zone of soil surrounding the root that is subject to influence by the root. Rhizobacteria are plant-associated bacteria that are able to colonize and persist on roots. An understanding of the ecology of a microorganism is a fundamental requirement for the introduction of a microbial inoculant into the open environment. This is particularly true for biological control of root pathogens in the rhizosphere, where one is actively seeking to alter the ecological balance so as to favour growth of the host plant and to curtail the development of pathogens. Some strains of plant growth-promoting rhizobacteria can effectively colonize plant roots and protect plants from diseases caused by a variety of root pathogens and growth promotion of plants through direct stimulation of growth hormone. Such beneficial or plant health-promoting strains are emerging as promising biocontrol agents. They are suitable as soil inoculants either individually or in combination and may be compatible with current chemical pesticides. Considerable progress has been achieved using molecular genetic techniques to elucidate the important microbial factors or genetic traits involved in the suppression of fungal root diseases. Strategies utilizing molecular genetic techniques have been developed to complement the ongoing research ranging from the characterization and genetic improvement of a selected biocontrol agent to the measurement of its persistence and dispersal. Finally, biocontrol is considered as part of a disease control strategy like integrated pest management which offers a successful approach for the deployment of both agro-chemicals and biocontrol agents.     

Using evolutionary tools to facilitate the prediction and prevention of host‐based differentiation in biological control: a review and perspective

The unprecedented success of biological control (biocontrol) agents led some of the proponents of this technology to promote its use as a panacea for all pest problems. Following an accumulation of non‐target host interactions, because of generalist or new association introductions, techniques to help ensure classical biocontrol agent's success and reduce non‐target interactions were implemented. Even with these new measures in place, public and scientific mistrust and lack of consistency has resulted in increased regulation of biocontrol introductions. This has likely decreased the probability of effective, sustainable control measures being expeditiously implemented. With the current apprehension concerning the safety of biocontrol, we should incorporate the processes (adaptation, selection, etc.) and theoretical concepts of evolutionary biology to predict and enhance the effectiveness of biocontrol. The microevolutionary perspective that involves mutation, drift, selection and gene flow may be a crucial consideration in the realm of biocontrol. Here, we discuss how and why spatial and evolutionary models should be implemented into future risk assessment analyses of potential biocontrol agents. We suggest that it is necessary to re‐assess the approach that has developed over the past approximately 100 years of sustained releases and illuminate them in the context of an evolutionary timescale.     

Wood Chip-Polyacrylamide Medium for Biocontrol Bacteria Decreases Verticillium dahliae Infection on Potato

The lack of consistent success of biological control of soilborne plant pathogens may be due to the introduction of the organism into a foreign environment. We hypothesized that wood chip-polyacrylamide (PAM) cores surrounding host plant roots could alter the soil environment to favour growth of introduced biocontrol microorganisms, thereby reducing Verticillium dahliae infection of potato ( Solanum tuberosum L.) in a greenhouse. A 7cm diameter ×15cm deep hole (core) was drilled in the center of a 20 ×30cm deep pot (1.9 kg) containing soil infested with V. dahliae inoculum. Cores were then filled with wood chip-PAM-biocontrol organism mixtures. Soils that had Streptomyces lydicus inoculated into wood chip-PAM cores had lower levels of V. dahliae symptoms ( V vis ) and V. dahliae isolations ( V iso ) than all other treatments in three soils. V vis and V iso on plants growing in soils amended with S. lydicus or Pseudomona corrugata inoculated into the soil itself (without wood chip-PAM cores) did not differ from soils that were unamended with these biocontrol organisms. V. dahliae biomass was lower in wood chip-PAM cores inoculated with S. lydicus than control or wood chip-PAM cores without biocontrol bacteria. Soils with wood chip-PAM cores inoculated with S. lydicus or P. corrugata generally had higher microbial biomass/ V. dahliae biomass (MB/VB) ratios than control soils, or soils with S. lydicus or P. corrugata inoculated into the soil. Wood chipPAM cores alone and wood chip-PAM cores inoculated with S. lydicus had higher MB/VB ratios than wood chip-PAM cores inoculated with P. corrugata . V vis and V iso were curvilinearly correlated with the MB/VB ratios in negative relationships, respectively (r 2 = 0.68, r 2 = 0.68). As the MB/VB ratio increased, V vis and V iso decreased. Although field studies and economic evaluations are necessary, amending soil with wood chips-PAM and a biocontrol bacterium may be a valuable method to increase the effectiveness of biocontrol organisms.     

Chitinolytic activity of endophytic Streptomyces and potential for biocontrol

Aims: Biological sources for the control of plant pathogenic fungi remain an important objective for sustainable agricultural practices. Actinomycetes are used extensively in the pharmaceutical industry and agriculture owing to their great diversity in enzyme production. In the present study, therefore, we evaluated chitinase production by endophytic actinomycetes and the potential of this for control of phytopathogenic fungi. Methods and Results: Endophytic Streptomyces were grown on minimum medium supplemented with chitin, and chitinase production was quantified. The strains were screened for any activity towards phytopathogenic fungi and oomycetes by a dual‐culture in vitro assay. The correlation between chitinase production and pathogen inhibition was calculated and further confirmed on Colletotrichum sublineolum cell walls by scanning electron microscopy. Conclusions: This paper reports a genetic correlation between chitinase production and the biocontrol potential of endophytic actinomycetes in an antagonistic interaction with different phytopathogens, suggesting that this control could occur inside the host plant. Significance and Impact of the Study: A genetic correlation between chitinase production and pathogen inhibition was demonstrated. Our results provide an enhanced understanding of endophytic Streptomyces and its potential as a biocontrol agent. The implications and applications of these data for biocontrol are discussed.     

Suppression of Fusarium wilt by combining green compost and Trichoderma hamatum

Fusarium wilts, caused by the fungus Fusarium oxysporum, are important diseases of horticultural and agricultural crops and lead to significant yield losses. The pathogen infects the roots and colonizes the vascular tissue, leading to wilting and finally death of the plant. The objective of this study was to investigate the efficacy of amendment of green compost and a Trichoderma hamatum strain against Fusarium wilt of radish. The substrate effects and the effect of a Trichoderma strain were tested in a potting soil bioassay. The tested composts lowered the disease level and had a positive influence on the plant yield (fresh weight and dry weight). Nothwithstanding, only a small dosis effect of the amendment was observed. In the presence of the tested Trichoderma hamatum strain no significant lower disease level was observed. Furthermore, a minor effect on plant yield (fresh weight and dry weight) was observed compared to the amendment with only green compost. Maybe the pathogenic Fusarium oxysporum strain and the Trichoderma strain competed for nutrients, iron could be a limiting factor. One possible approach to improve biological control may be the application of combinations of biocontrol agents.     

Rate of consumption of Western flower thrips pupae by the soil-dwelling mite Hypoaspis sclerotarsa (Acari: Laelapidae)

Western flower thrips (WFT), Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), are one of the most serious pests of horticultural crops worldwide. Despite one third of its lifecycle being in the soil, the potential for biological control of WFT in the soil is poorly understood and requires further elucidation. A number of studies report that polyphagous predatory mites prey on pupal stages of WFT in the soil, but little has been done on consumption rates. Therefore, we designed a laboratory study to examine the rate of consumption of WFT pupae in potting media, by the soil-dwelling mite Hypoaspis sclerotarsa (Costa). Five predator densities were evaluated (0, 2, 4, 6 and 8) against four densities of WFT prey (5, 10, 15 and 20 pupae). Pupal consumption was assessed at 2 hourly intervals over a six-hour period. The study confirmed that H. sclerotarsa did consume WFT pupae and that the rate of consumption increased with increasing density of H. sclerotarsa. The rate of consumption also increased with the density of WFT pupae. However, this was not consistent because, as the numbers of WFT pupae increased, so did the ratio of WFT pupae remaining to those consumed, increase. This paper is the first report of H. sclerotarsa in Kenya, and of its potential as a biocontrol agent of WFT. Further studies are now needed to understand interaction of foliar and soil dwelling predatory mites (H. sclerotarsa) for control of WFT under field conditions.     

Success and safety in the biological control of environmental weeds in New Zealand

Biological control, using specialist insect herbivores and plant pathogens, can be a self‐sustaining, cost‐effective and low‐risk tool for the management of environmental weeds. Agents have been recorded attacking non‐target plants in New Zealand and elsewhere, but the effects are usually minor and/or transitory. It seems probable that only two cases, worldwide, will result in significant damage to non‐target plants (representing 0.5% of the nearly 400 insect, mite, or fungal species used in classical weed biocontrol). Both of these cases were predictable from host range testing. Negative indirect, or ‘downstream’, ecological effects from specific weed biocontrol agents are difficult to predict and measure. They are probably insignificant compared to the impacts of the invasive plants that the agents are introduced to control. However, it is necessary to balance the risks associated with any introduction against the environmental benefits from controlling a weed to a predicted level. Recent analyses suggest that success rates are better than generally perceived. For New Zealand programmes, where enough time has lapsed to allow assessment, we calculate a full/partial success rate of 83%. Many of the costs associated with environmental weeds are difficult to quantify. Detailed risk assessment will make biological control programmes more expensive and time‐consuming, so that reliance on non‐biological management methods for environmental weeds may actually increase. The costs of biocontrol programmes against some New Zealand weeds can be kept down by using research already carried out in Australia and other countries, and the process is reciprocal. Developing international consortia of sponsors is also a potential way to fund programmes against weeds shared by several countries.     

Multiple pathways for invasion of anurans on a Pacific island

Since 1937, thirteen species of non-indigenous anurans have made their way to Guam. Of these, at least six have established breeding populations. Various pathways led to the introduction of these species to the island. The only anuran intentionally introduced was Chaunus marinus (formerly Bufo marinus ), which was brought to Guam as a biocontrol agent. Kaloula picta, K. pulchra, Polypedates leucomystax , and probably Litoria fallax arrived as stowaways via maritime or air-transport vessels. Eleutherodactylus coqui and Euhyas (formerly Eleutherodactylus ) planirostris appear to have entered Guam through the horticultural trade. Specimens of Pseudacris regilla were found among agricultural products and Christmas trees. Five species have been transported to Guam via the aquacultural trade. The importation of tilapia, milkfish, and white shrimp from China, Hong Kong, Taiwan, and the Philippines was associated with the introduction to Guam of Fejervarya cancrivora, F. limnocharis sensu lato, Microhyla pulchra, Polypedates megacephalus , and Sylvirana guentheri (formerly Rana guentheri ). Presently, no quarantine or containment guidelines have been established for Guam's aquacultural industry.     

Selection responses and quantitative-genetic analysis of preadult performance on two host plants in the bean weevil, Acanthoscelides obtectus

The infectivity and biocontrol potential of entomopathogenic nematodes against two common urban tree leaf beetles (Altica quercetorum and Agelastica alni) pupating in the soil were examined under laboratory and semi‐field conditions. In the laboratory experiments, pre‐pupae and pupae of both insect species were shown to be highly susceptible to nematode infection when challenged in soil pre‐treated with the parasites’ infective juveniles. In general, Heterorhabditis megidis was more effective than Steinernema feltiae. However, significant differences were observed between individual isolates within the latter species. Nematodes developed and reproduced in cadavers of both insect species. A semi‐field experiment studying the biocontrol potential of selected nematode strains, conducted under the canopy of urban trees, confirmed the preliminary laboratory findings and revealed that H. megidis could eliminate most of the insects pupating in the soil, when applied at a relatively low dose of 10⁵ IJs m^?2. The potential of entomopathogenic nematodes as environmentally safe, effective, and economically viable agents for the biological control of tree leaf beetles in urban green areas is discussed.     

Multicellular behaviour and production of a wide variety of toxic substances support usage of Bacillus subtilis as a powerful biocontrol agent

Intensive cultivation of plants in the monoculture field system in order to feed the continuously growing human population creates a need for their protection from the variety of natural competitors such as: bacteria, fungi, insects as well as other plants. The increase in the use of chemical substances in the 20th century has brought many effective solutions for the agriculture. However, it was extremely difficult to obtain a substance, which would be directed solely against a specific plant pathogen and would not be harmful for the environment. In the late 1900's scientists began trying to use natural antagonisms between resident soil organism to protect plants. This phenomenon was named biocontrol. Biological control of plants by microorganisms is a very promising alternative to an extended use of pesticides, which are often expensive and accumulate in plants or soil, having adverse effects on humans. Nonpathogenic soil bacteria living in association with roots of higher plants enhance their adaptive potential and, moreover, they can be beneficial for their growth. Here, we present the current status of the use of Bacillus subtilis in biocontrol. This prevalent inhabitant of soil is widely recognized as a powerful biocontrol agent. Naturally present in the immediate vicinity of plant roots, B. subtilis is able to maintain stable contact with higher plants and promote their growth. In addition, due to its broad host range, its ability to form endospores and produce different biologically active compounds with a broad spectrum of activity, B. subtilis as well as other Bacilli are potentially useful as biocontrol agents.