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In plant pathology, the term applies to the use of microbial antagonists to suppress diseases as well as the use of host-specific pathogens to control weed populations. In both fields, the organism that suppresses the pest or pathogen is referred to as the biological control agent (BCA). More broadly, the term biological control also has been applied to the use of the natural products extracted or fermented from various sources. These formulations may be very simple mixtures of natural ingredients with specific activities or complex mixtures with multiple effects on the host as well as the target pest or pathogen. And, while such inputs may mimic the activities of living organisms, non-living inputs should more properly be referred to as biopesticides or biofertilizers, depending on the primary benefit provided to the host plant. The various definitions offered in the scientific literature have sometimes caused confusion and controversy. For example, members of the U.S. National Research Council took into account modern biotechnological developments and referred to biological control as “the use of natural or modified organisms, genes, or gene products, to reduce the effects of undesirable organisms and to favor desirable organisms such as crops, beneficial insects, and microorganisms”, but this definition spurred much subsequent debate and it was frequently considered too broad by many scientists who worked in the field (US Congress, 1995). Because the term biological control can refer to a spectrum of ideas, it is important to stipulate the breadth of the term when it is applied to the review of any particular work.
In contrast, antagonism between organisms results in a negative outcome for one or both. Competition within and between species results in decreased growth, activity and/or fecundity of the interacting organisms. Biocontrol can occur when non-pathogens compete with pathogens for nutrients in and around the host plant. Direct interactions that benefit one population at the expense of another also affect our understanding of biological control.
Significant biological control, as defined above, most generally arises from manipulating mutualisms between microbes and their plant hosts or from manipulating antagonisms between microbes and pathogens.
Stimulation of plant host defense pathways by non-pathogenic BCAs is the most indirect form of antagonism. However, in the context of the natural environment, most described mechanisms of pathogen suppression will be modulated by the relative occurrence of other organisms in addition to the pathogen. While many investigations have attempted to establish the importance of specific mechanisms of biocontrol to particular pathosystems, all of the mechanisms described below are likely to be operating to some extent in all natural and managed ecosystems. And, the most effective BCAs studied to date appear to antagonize pathogens using multiple mechanisms.
Because plant diseases may be suppressed by the activities of one or more plant-associated microbes, researchers have attempted to characterize the organisms involved in biological control. Historically, this has been done primarily through isolation, characterization, and application of individual organisms. By design, this approach focuses on specific forms of disease suppression. Specific suppression results from the activities of one or just a few microbial antagonists. This type of suppression is thought to be occurring when inoculation of a biocontrol agent results in substantial levels of disease suppressiveness. Its occurrence in natural systems may also occur from time to time. For example, the introduction of Pseudomonas fluorescens that produce the antibiotic 2,4-diacetylphloroglucinol can result in the suppression of various soilborne pathogens (Weller et al. 2002). However, specific agents must compete with other soil- and root-associated microbes to survive, propagate, and express their antagonistic potential during those times when the targeted pathogens pose an active threat to plant health. In contrast, general suppression is more frequently invoked to explain the reduced incidence or severity of plant diseases because the activities of multiple organisms can contribute to a reduction in disease pressure. High soil organic matter supports a large and diverse mass of microbes resulting in the availability of fewer ecological niches for which a pathogen competes. The extent of general suppression will vary substantially depending on the quantity and quality of organic matter present in a soil (Hoitink and Boehm 1999). Functional redundancy within different microbial communities allows for rapid depletion of the available soil nutrient pool under a large variety of conditions, before the pathogens can utilize them to proliferate and cause disease. For example, diverse seed-colonizing bacteria can consume nutrients that are released into the soil during germination thereby suppressing pathogen germination and growth (McKellar and Nelson 2003). Manipulation of agricultural systems, through additions of composts, green manures and cover crops is aimed at improving endogenous levels of general suppression.
Most pathogens will be susceptible to one or more biocontrol strategies, but practical implementation on a commercial scale has been constrained by a number of factors. Cost, convenience, efficacy, and reliability of biological controls are important considerations, but only in relation to the alternative disease control strategies. Cultural practices (e.g. good sanitation, soil preparation, and water management) and host resistance can go a long way towards controlling many diseases, so biocontrol should be applied only when such agronomic practices are insufficient for effective disease control.
In general, though, regulatory and cultural concerns about the health and safety of specific classes of pesticides are the primary economic drivers promoting the adoption of biological control strategies in urban and rural landscapes.
In terms of efficacy and reliability, the greatest successes in biological control have been achieved in situations where environmental conditions are most controlled or predictable and where biocontrol agents can preemptively colonize the infection court. Monocyclic, soilborne and post-harvest diseases have been controlled effectively by biological control agents that act as bioprotectants (i.e. preventing infections). Specific applications for high value crops targeting specific diseases (e.g. fireblight, downy mildew, and several nematode diseases) have also been adopted. As research unravels the various conditions needed for successful biocontrol of different diseases, the adoption of BCAs in IPM systems is bound to increase in the years ahead.
Trichoderma harzianum • Trichoderma virens
mixed path antagonism
Chitinases • Glucanases • Proteases
Compost Teas • Exudates • leachates • Physical niche occupation