Neil Aldrin Valeroso

II GLA / SSP B

Ecological backlash and its management

Ecological backlash involves the counter-responses of pest populations or other biotic factors in the environment that diminish the effectiveness of pest management tactics. Many of these counter responses result from heavy mortality burdens imposed by the pest management tactics. Other responses arise from disruption of ecological processes or changes in resource levels of the biotic community. Ecological backlash mainly manifests in the form of resistance, resurgence, and replacement- the three "R" s of pest management awareness. In this lecture the following aspects of ecological backlash will be discussed: Resistance of populations to management tactics - principles of resistance - resistance to conventional insecticides, mechanisms of resistance, cross-resistance. - resistance to other management tactics - management of resistance Pest population resurgence and replacement - causes for resurgence and replacements - managing resurgence and replacement Other forms of ecological backlash

Resistance of populations to pest management tactics

Resistance is defined as the ability of certain individuals to tolerate or avoid a dose that would prove lethal or reproductively degrading to majority of individuals in a normal population. Roman Sawicki (1987) proposed an improved definition: "Resistance is a genetic change in response to selection by toxicants that may impair control in the field".

Principles of resistance Resistance in a population can be explained by the same principles that explain evolution by natural selection. Natural selection or darwinian selection, is a selection of the fittest genotype for a given set of environmental conditions. Selection for pesticide resistant genotypes is a similar process. The rate of resistance development i.e. time required for the development of resistance depends on - mortality burden imposed on the population by the management tactic - genetic factors; Monogenic and polygenic resistance Resistance to conventional insecticides Resistance to insecticides is the most preeminent form of resistance in insects. The prevalence of this phenomenon reflects the widespread effectiveness of these compounds in causing pest mortality. Magnitude of the insecticide resistance problem: The first widely recognized case of pesticide resistance among agricultural arthropod pests was reported in the early 1900s, when lime sulfur failed to control the San Jose scale, Quadraspidiotus perniciosus, in Washington State. In 1946, house flies were discovered resistant to DDT in Sweden. Since then, more than 500 species of arthropod pests are known to have evolved resistance to one or more major groups of insecticides, including organochlorines, organo phosphates (OPs), carbamates, and synthetic pyrethroids. This total includes 375 cases or 75% among agricultural insects and 125 or 25% among pests of medical importance to humans or animals. Resistance in 31 species of predators and parasitoid species has also been documented. At least 17 species have developed resistance to all major groups of insecticides. Mechanisms of resistance to insecticides Insects have evolved three major mechanisms to overcome toxicants; 1) biochemical resistance; 2) physiological resistance; and 3) behavioral resistance.

1. Biochemical resistance: In this form of resistance, an insecticide is detoxified by one or more enzymes before it can reach its site of action. Mixed-function oxidases or other enzymes are involved. 2. Physiological resistance: Physiological resistance is any form of resistance that reduces toxicity through changes in basic physiology. In this form of resistance, the chemical is not broken down into a less toxic form, rather the insect accommodates the chemical by altering one or more physiological functions. This may involve - reduced neuronal sensitivity to insecticides e.g., knockdown resistance (kdr) in house flies to DDT and pyrethroids. - altered acetylcholinesterase. Acetylcholinesterase that is less sensitive to inhibition by OP and carbamate insecticides has been documented in resistant strains of several insect, tick and mite species. E.g. Anopheles mosquito s resistant to OP and carbamates have been shown to have an altered acetylcholinesterase. - decreased penetration of the insecticide through the body wall through modification of the structure or composition of the cuticle. e.g., additional waxy layers in resistant strains. - increased excretion or sequestration of insecticide. e.g. DDT storage in body fat, preventing it from reaching the site of action. 3. Behavioral resistance: involves changes in behavior by which insects avoid insecticides. Ex. In mosquito, Anopheles gambiae, an endophillic strain (indoor dwelling) was susceptible to DDT sprays applied to the indoor walls. An exophillic strain not inhabiting indoors became dominant because its behavior allowed it to avoid exposure to the insecticide. Cross-resistance: ability of an insect with resistance to one insecticide to resist other insecticides. Cross-resistance necessitates changing from one insecticide to another to obtain pest suppression. Colorado potato beetle, Leptinotarsa decemlineata, has developed resistance to - arsenicals in the 1940s - chlorinated hydrocarbons, OPs, and carbamates in 1970s - synthetic pyrethroids in 1980s - currently the chloronicotinyl insecticide, imidacloprid, is most effective.

Cross-resistance to insecticides can be within a class of insecticides or between classes with similar modes of action. House flies resistant to knockdown by DDT are also resistant to pyrethroids. This resistance is conferred by the recessive kdr gene. House flies which are cross-resistant to OPs and carbamates through altered acetylcholinesterase, receive this trait through inheritance of the codominant AChE-R gene. Resistance to other pest management tactics 1. Resistance to insect growth regulators (IGRs): IGRs disrupt development processes of insects and other arthropods. Cross-resistance in various insecticide-resistant strains has been a major factor in IGR resistance. Currently, at least 13 insect species representing Diptera, Coleoptera, Homoptera, and Lepidoptera show cross-resistance to IGRs. Resistance has been documented so far to methoprene, hydroprene, kinoprene, and diflubenzuron. Resistance appears to be due to reduced penetration and increased metabolism of the compounds. 2. Resistance to microbial insecticides: Resistance has been reported against bacterial and viral insecticides. Resistance to Bacillus thuringiensis products has been reported in Indianmeal moth, Plodia interpunctella, diamondback moth, Plutella xylostella. Resistance to viral insecticides have been observed in several Lepidoptera and Hymenoptera. E.g., reported resistance in spruce budworm to viral insecticides. 3. Resistance to plant rotations: This form of resistance has been documented in northern corn rootworm, Diabrotica barberi, a major pest of corn. 4. Virulence to resistant plants: We already talked about biotype formation in the last class. The occurrence of insect biotypes has been documented in several species of aphids, hessian fly, and brown planthopper. In addition to resistance development to the above pest management tactics, there are reports of resistance development to parasites and to sterile-male release technique in screwworm fly.

Management of resistance In this section, we will mainly consider resistance management to insecticides because of its widespread occurrence. However, the same principles of insecticide management used to slow resistant development apply generally to other types of resistance. Conditions that promote resistance: The basic cause of resistance development is the high mortality (or reduced reproduction) rate placed on the insect population, resulting in an inadvertent selection for individuals capable of overcoming the burden. Some of the factors that influence the rate of resistance development are as follows: Operational factors: 1) Exposure time to a single insecticide- longer the exposure time to a single insecticide or to a single slow-release insecticide, greater the rate of resistance development 2) Number of generations of the insect exposed- is insecticide applied against every generation? 3) High mortality burden- high selection pressure 4) Coverage by the insecticide- are any parts of the crop not covered by the sprays 5) Geographical area covered- are all populations in an area are treated? 6) The insecticide is closely related to the one used before 7) Low economic threshold is employed 8) Insecticide is inherently irritating and/or repellent Biological factors: 1) Extent of migration 2) Species is a monophagous insect 3) Short generation time 4) Numerous offspring per generation are produced 5) Highly mobile species- greater probability of exposure Slowing the process of resistance development: Resistance development inevitable if mortality burdens are high.

Rate of resistance development can be slowed by considering the operational factors that enhance it and modifying the management tactics accordingly. Integration of several tactics Passive tactics Other methods suggested to disrupt or slow resistance development include: A. Management by moderation: Some of the elements involved in this approach are as follows: 1) Use of low doses of insecticides, preserving some of the susceptible genotypes. 2) Less frequent insecticide applications 3) Use chemical with short residual activity 4) Avoid slow release formulations 5) Apply selection against adults after reproduction 6) Make local applications 7) Leave some generations or populations untreated 8) Treat only parts of the crop 9) Use higher economic thresholds B. Management by saturation: this is accomplished by the following tactics: 1. Render the resistance gene "functionally" recessive with extremely high doses of the insecticide 2. Suppress detoxification mechanisms with the use of synergists: Synergists function by inhibiting specific detoxification enzymes. For example, piperonyl butoxide is an oxidase inhibitor, and is commonly added to insecticides to enhance suppression at low to moderate levels of resistance. C. Management by multiple attack: This approach is very similar to the integrated system discussed earlier, except that only insecticides are employed here. 1. Use insecticide mixtures: mechanisms of resistance are different for each member chemical, and that the mechanisms required to overcome resistance are not present in any one individual. Mixture of temephos, propoxur, and permethrin for the control of mosquito s.

2. Apply insecticides in a mosaic pattern across an area: unrelated chemicals are applied in different parts of the pest population range. Insects not killed by one chemical in a sector will get killed when they move into another sector treated with a different chemical. E.g. opposite walls of a house are treated with different chemicals to suppress horn flies. 3. Apply insecticides in a rotation scheme: use of dissimilar insecticides in a rotation will result in reduced frequency of resistant individuals between the times the "resistant " chemical is used.

Pest population resurgence and replacement

Resurgence: is defined as a situation where a population, after having been suppressed , rebounds to numbers greater than before suppression occurred. Replacement: also referred as secondary pest outbreak, occurs when a major pest is suppressed and is continued to be suppressed by a pest management tactic, but is replaced by another pest previously with minor pest status. During secondary pest outbreaks, the tactic has an effect on the primary pest, but has no effect on the secondary pest. Three major causes have been suggested to explain resurgence and replacement. 1) Reduction of natural enemies along with the pest 2) Direct favorable influences on physiology and behavior of pests 3) Removal of competitive species

1) Upsets from reduction of natural enemies: Mortality in natural enemies is caused by direct exposure to toxicants and also indirectly due to a) starvation as a result of host removal, b) death from secondary poisoning due to feeding on contaminated prey, and c) repellency of the insecticide. Paradigm for resurgence: Populations of both natural enemies and the pest species are reduced simultaneously as a result of insecticide application. As the residual effects of the insecticide wane, the pest population grows, but because the natural enemies must wait for adequate food supplies, their populations lag in regrowth. Unencumbered by natural enemies, pest population growth accelerates and growth exceeds previous levels.

An example of resurgence: Cyclamen mite, Steneotarsonemus pallidus, and its mite predator, Tephlodromus reticulatus system on strawberries in California. Paradigm for pest replacement: Insecticide application reduces the populations of the key pest; at the same time natural enemies of the secondary pests are also destroyed, but causes little mortality of the secondary pest itself. Lacking the natural enemy check, the secondary pest population multiplies and attains a key pest status. Example: Michigan apple system - spider mite resurgence. 2) Favorable effects of pesticides on arthropod physiology and behavior: In many instances, sublethal doses of pesticides have been shown to have favorable effects on arthropod physiology and/or behavior. Luckey in 1968 proposed the term "hormoligosis" (from the Greek hormo = excite, oligo = small quantities) to describe this phenomenon. Luckey provided data to support that sublethal doses of several insecticides increased the growth rate of the cricket, Acheta domesticus. 3) Upsets from removal of competitors: If two or more species are competing in an area for the same requisite and one species is dominant, removal of the dominant species may result in replacement by the subordinate species, irrespective of changes in natural enemy populations. Managing resurgence and replacement The fundamental objective of programs designed to reduce resurgence and replacement is to avoid hormoligosis and reduce destruction of natural enemies in the agroecosystem. Another approach is to restore natural enemy populations through inoculative releases. Avoiding hormoligosis: Follow all specifications on the label, apply on target, and minimize drift. Avoiding natural enemy destruction: Most of the approaches are based on insecticide selectivity. Selectivity is the use of pesticides to kill plant-feeding arthropods while not affecting their entomophagous natural enemies, and other non target species. Insecticide selectivity can be achieved in one of the following two ways. First approach is to use selective chemicals that have stronger suppressive effects on the pest population than on its natural enemies. This form of selectivity is referred to as physiological selectivity. Another approach is to use nonselective chemicals in a selective manner. This form of selectivity has been called ecological selectivity.

Physiological selectivity: Most conventional insecticides, with few exceptions such as trichlorfon, pirimicarb, and certain miticides, are nonselective. Some of the more important examples of physiological selectivity are found with nonconventional materials like, microbial insecticides like Bts, insect growth regulators, nonsteroidal ecdysone mimics like tebufenozide etc. Pyrethroids also may offer a degree of selectivity with certain pest-natural enemy systems. Ecological selectivity: Major approaches to ecological selectivity are as follows: 1) Use economic threshold levels 2) Treat with lowest rates possible 3) Avoid treating broad areas 4) Time treatments to avoid natural enemy destruction 5) Use the most selective formulation Inoculative releases of natural enemies: Another management approach that has been used to prevent pest upsets is to repopulate an area with inoculative releases of natural enemies after insecticide application. This tactic involves application of insecticides when pest populations cannot be regulated by natural enemies, realizing that natural enemy population will also be destroyed. Insectary reared natural enemies are made after insecticide residues have subsided to enhance natural control. Release of insectary reared resistant natural enemiesOther forms of ecological backlash Enhanced microbial degradation: This phenomenon has been documented against carbofuron applied to control corn rootworm in corn fields. Upsets in community balance: Community upsets occur when we change status quo by applying pest suppression tactics. Ex. Screwworm fly-White-tailed deer-tick parasite.