ReviewImproving the efficacy of biocontrol agents against soilborne pathogens☆
Introduction
Meeting the goal of improving soilborne disease control for efficient and sustainable production systems requires a reduction of the chemical inputs in agriculture. Various movements or parties see increased pesticide application as a threat to public health and the environment. Agronomists experience growing problems with the buildup of resistance to the pesticides in target pathogen populations (Dekker, 1976; Brent and Hollomon, 1998) and the withdrawal of pesticides from the market (Gullino and Kuijpers, 1994). Of special relevance, in the case of many vegetable and ornamental crops, is the loss of methyl bromide as soil fumigant (Katan, 1999; Gullino et al., 2003). Furthermore, the private industry is coming across increasing costs in discovering and developing new molecules for disease control application. Consequently, there is a growing awareness that integrated pest management (IPM) strategies can provide more environmentally sound and economically feasible alternatives for soilborne disease control. Biological control with introduced microorganisms has been identified as an important component of IPM strategies (Cook, 1993).
Many studies have been devoted to the identification of microorganisms able to reduce the activity of soilborne pathogens during the past four decades (Baker and Snyder, 1965).
Many laboratories around the world have developed their own microorganisms and this allowed the collection of important contributions about the biology of pathogens and antagonists. However, in spite of the initial great optimism and extensive later research efforts, progress in achieving commercial, large-scale usage of biological pest control has been slow.
Most experiences were on a laboratory scale, thus avoiding the problems related to antagonist mass production and biofungicide formulation, and disease control trials were performed in simplified environments such as growth chambers, experimental greenhouses or small field plots, thus avoiding the risk of large-scale experiments. When trials move towards the farm scale, many antagonists fail to confirm their activity or, when tested in different environments, behave unpredictably, and consequently lack reliability (Mathre et al., 1999).
Biofungicides presently meet many constraints and it is not a straightforward task to find, develop and implement practically feasible biocontrol products against soilborne diseases. Nevertheless, there are also reasons for stating careful optimism. There are many possibilities for combining various biocontrol agents, with each other, or with agronomical, physical or chemical control methods. In particular, by combining different methods of control, the aim is to obtain a synergistic effect, rather than additive. For that reason, a complete comprehension of the mode of action and mechanism of control is needed. A good combination is obtained when both partners benefit from the combined treatment. For example, combining a biocontrol agent (BCA) with a fungicide improves the control by the BCA and, at the same time, it enables to reduce the fungicide dosage, rendering its use more acceptable. Moreover, the combination of control methods provides a wider spectrum of control, which is especially needed to replace the wide-spectrum fumigants, such as methyl bromide. Finally, combining different control methods enables the use of less effective control methods which cannot stand by themselves, but in combination they can contribute to the control.
Other possibilities to improve the antagonist effectiveness include the genetic manipulation of the microorganisms and the enhancement of the mass production, formulation and methods of application The main possibilities to improve the efficacy of the biocontrol agents against soilborne pathogens are the topic of this brief review.
Section snippets
Combination with chemical pesticides
The combined use of BCAs (biocontrol agents) and chemical pesticides has attracted much attention in order to obtain synergistic or additive effects against the target organisms (Locke et al., 1985). Reduced amounts of fungicide can stress and weaken the pathogen and render its propagules more susceptible to subsequent attack by the antagonist (Hjeljord and Tronsmo, 1998). Replacement of some of the chemical fungicide treatments with BCAs does not only reduce the input of chemicals but can also
Combination with soil disinfestation
Soil disinfestation is commonly used for sanitation in low-technology houses and fumigation with chemicals, heating, steaming and solarization are the main methods in use (Garibaldi and Gullino, 1995). Soilborne pathogens were effectively controlled by using methyl bromide (Bell et al., 1996). However, concern regarding the potential of methyl bromide to deplete ozone led to its inclusion among the substances controlled by the Montreal protocol whose use must be eliminated by the end of 2004
Application of agronomical practices for biocontrol
Several agronomical methods permit to promote biocontrol agents. These include advantageous regimes for choosing crop plants, soil types, crop rotations, soil amendments, and the application of suitable ploughing, planting and sowing regimes.
Natural disease suppressive soils are good examples in which the indigenous microflora effectively protects plants against soilborne pathogens. Suppressive soils have been described for many soilborne pathogens, including Fusarium oxysporum (Scher and
Mixtures of antagonists
Association of several microorganisms is needed to control different diseases that affect the same crop. Most of the BCAs are specific only for a given type of target pathogen. Although this property represents an advantage from the environmental point of view, it creates great difficulties to the growers who need to control several plant pathogens in the same crop. Moreover, the combination of two or more antagonists means also multiple registration processes, with increased costs and
Genetic manipulation
Despite years of research and development, significant questions regarding the physiological and ecological constraints that limit biological control remain unanswered. Molecular and genomic tools offer new possibilities for improving the selection, characterization, and management of biological control. Development in proteomics and functional genomics will give us the possibility to determine and follow expression of crucial genes of BCA's during mass production, formulation, storage and
Mass production
A critical factor that must be considered when selecting a BCA for commercial development is the availability of a cost-effective production and stabilization technology that yields an optimally effective form of the antagonist. More studies on the practical aspects of mass production and formulation need to be undertaken to make new biocontrol products stable, effective, safer and more cost effective (Fravel et al., 1999).
Major characteristics to market a biofungicide are the following (Agosin
Formulation and methods of application
Biological control has often been idealized as a method for controlling plant diseases, as has happened for organic farming compared to the traditional cropping techniques. In organic farming, the bottleneck is represented by keeping remunerative yield, lowering the selling price of the product. For biological control, the major difficulty to reach the market and to be competitive with the chemical fungicides is represented by a consistent and reliable effectiveness and by the length of shelf
Conclusions
The history of biological control of plant pathogens is not so recent. Sandford in 1926 started biocontrol against potato scab, but research on this topic received a renewed impetus and attracted many scientists after the 1963 International Symposium held at Berkeley on ecology of soilborne plant pathogens: prelude to biological control (Baker and Snyder, 1965; Baker and Cook, 1974). Despite many decades of research in biological control, biopesticides represent about 1% of the global pesticide
Acknowledgements
Work carried out with a grant from the Italian Ministry for the Environment and Territory within the Framework Agreement “Sustainable Agriculture”.
The authors thank Professors Jaakov Katan and Angelo Garibaldi for their critical readings.
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Invited paper at the 15th International Plant Protection Congress, Beijing, China, 11–16 May 2004.