The usual agroecological practices for preventing soilborne disease are well known: crop rotation, addition of organic matter and compost, sanitation of tools and plant materials, promotion of good soil structure, prudent water management, use of resistant varieties. Add to that list an incongruous entry — continuous monoculture. When the same crop gets hammered by the same disease in the same field year after year, once in a while the disease will suddenly disappear for good. This is the phenomenon of suppressive soil, and Daniel Schlatter and colleagues recently summarized the latest research on it.
While continuous monocropping contravenes good agroecological management on many levels, the appearance of disease suppression actually does arise out of agroecological principles. What we call disease is really the interaction between a host, in this case a crop plant, and a pathogenic organism. And whenever an organism is present in a field, it is subject to ecological forces. A field that contains a large population of a single species, whether it be a plant, insect, or microbe, potentially presents a target for another organism. For a plant, the opportunistic organism could be an insect or a fungus. For a microbe responsible for a crop disease, the opportunist could be an antagonist — a sort of armed competitor — or even a hyperparasite.
To be sure, a certain amount of disease suppression can come from enrichment of soil organic matter. Schlatter et al. differentiate this phenomenon as general suppression, the suppression that prudent organic farmers enjoy, whereby most soilborne diseases can cause only the most minimal damage. The other kind of suppression, that which arises from monocropping, is designated as specific suppression, where a specific disease caused by a single organism is the only one affected. Specific suppression can often be introduced into a field by addition of a small amount of already-suppressive soil. In addition, specific suppression requires the continuing presence of the host crop. If the field is rotated to a different crop, the resistance subsides, and it takes a season or more to return when the host crop is re-planted.
General suppression is thought to operate based on high numbers and high diversity of microbes present in the soil. This kind of microbial community insures that organic compounds are quickly broken down and taken up. Among these compounds are the exudates that a root characteristically gives off as it extends through the soil. It is these exudates that trigger pathogens to begin their attack. If the exudates dissipate immediately after release, the root may essentially slip past the pathogens unnoticed.
Regarding specific suppression, the authors of the review examine the mechanisms by presenting three case studies highlighting a continuum of suppression, from completely specific to somewhere between specific and general. The first regards the disease known as take-all of wheat. This disease is caused by a fungus that severely rots the plant’s roots. Certain strains of bacteria suppress the disease, and all of these turn out to produce an antibiotic known by the acronym DAPG. When these strains reach high population levels, the take-all pathogen is completely incapacitated and the disease disappears. The wheat plant’s roots even exude compounds that promote colonization by the DAPG strains. However, the DAPG producers have no effect on other pathogens.
The second case is that of Rhizoctonia bare patch of wheat and sugar beets. There is evidence that the organisms antagonistic to the Rhizoctonia fungus possess a certain amount of cross-antagonism against other pathogens, but the bare-patch-suppressing microbes are the least well characterized of the three case studies.
The third case is that of potato scab, a disease caused by a strain of Streptomyces, a genus of bacteria perhaps better known as the source of the antibiotic streptomycin. Interestingly, the antagonistic organisms are also strains of Streptomyces. It seems that the potato plant’s exudates are a food source for a suite of highly similar organisms, and niche theory predicts that in such a situation the best competitor will win out. In this case, the various strains of Streptomyces boost their competitive abilities in the presence of this limited food source by producing unique antibiotics that kill off their competitors. A casualty of this antibiotic arms race is the strain that causes the scab disease.
This third case of specific suppression is the closest to general suppression, although the mechanism is different from the fast degradation hypothesis. The cocktail of antibiotics pulsing through the soil can serve to eliminate other pathogens besides the bacterial strain responsible for potato scab. In fact, disease-suppressive Streptomyces strains can sometimes be enriched in the soil by incorporation of certain organic amendments, such as rice bran or the mustard family plant wall rocket, the presumption being that a particular type of food will cause the microbes that are able to digest it to fight over it, launching their specialized chemical weaponry as the competition intensifies. Whichever management practice initiates the chemical warfare, once it starts, it may tilt the microbial community composition toward ever stronger competitors, leading to a self re-enforcing dynamic that in some cases leads to lasting suppressiveness even when the original crop or amendment is gone.
Disease suppression has long been a phenomenon that emerged from the black box that is the soil microbiome. New tools for microbiome research are beginning to shed light on the inner workings of that system, and Schlatter and colleagues enumerate some of the methodological considerations for further advances. They also call for additional studies that use the knowledge that has been gained about suppression mechanisms to further our understanding of the agroecological circumstances and the specific management practices that will enhance disease suppression. For instance, will tillage increase or decrease disease-suppressive Streptomyces? Co-author Linda Kinkel invokes niche theory in predicting that no-till might allow the different strains to begin to coexist and to ease up on the competition, while tillage would disrupt any coexistence dynamic and more likely set off a chemical arms race. Would that happen in the real world? In every situation? Only field research, with a boost from microbiomics, will give the answer. For the foreseeable future, though, continuous monoculture remains a questionable bet for sustainable agriculture.
2 Replies to “the magic of monoculture”
Very interesting discussion of the difference between general and specific suppression. There is a general “theory” in IPM that a population of a pest has to build up to a certain level to be attractive enough to draw in effective biological control. I remember a colleague back in the 1970s who worked for United Fruit Co. managing its pest management programs in banana monocultures. He convince the company, at least for a time, to not worry so much when a pest appeared in a section of a large plantation, and predicted that the area of outbreak would come under control because of its attractiveness to control agents, be that microbial, entomological, or larger. His predictions worked so well that for a good number of years insecticides were not used on large areas of their production areas. Nervous farm managers after my colleague retired who were afraid to wait for the predators to catch up on the prey moved the company back into intensive insecticide application, and apparently more pest outbreaks. Is this the same thing going on in the microbiome of the soil? Does diversity replace this in agroecological systems?
It is akin to that IPM general theory, but a soil must also have suppressive potential for specific suppression to develop. Contrary to the environmental microbiologists’ quip, everything is NOT everywhere. The specific antagonist must be present in the soil, and other factors like soil type and microbial community may play a role. Regarding diversity, that is one of the factors required for general suppression, but general suppression operates by other mechanisms. An pest-regulation analogy for the fast degradation hypothesis of general suppression would be the apparency theory of plant defense, where diversity would mask the chemical and physical cues that allow a pest to find its host, although in this case the diversity would be made up of different vegetation rather than other arthropods. An analogy for the chemical arms race hypothesis might be the ant-plant mutualism, where a beneficial arthropod eliminates other arthropods on a host plant while receiving food from that very plant, although this analogy fails to capture the complex community dynamics of antibiotic-producing microbes.
As for one type of suppression replacing the other, the Holy Grail for suppressive soil researchers is to combine specific and general suppression in the same soil!