Both SIT and IIT can be combined together, and they are compatible with conventional biological control using parasitoid, predators, and pathogens. The release of Wolbachia -infected females may result in production of viable offsprings if the released females are compatible with either wild or released males [ 5 ].
Pesticides should only be used when necessary to keep pest populations below that cause economic damage. Selective pesticides, which have the least negative effects on the environment, should be used according to principles 5, 6, and 7 of IPM.
Botanicals and microbial biorational pesticides should be given priority in selection. The efficacy of these biorational pesticides may be increased when applied together [ 27 ].
The integration of a number of different control tactics into IPM systems can be done in ways that greatly facilitate the achievement of the goals either of field-by-field pest management, or of area-wide AW pest management, which is the management of the total pest population within a delimited area [ 1 ].
Knipling [ 45 ] used simple population models to demonstrate that small insect pest population left without management can compromise the efforts of containment of pest population in a large area. AW-IPM programs should be coordinated by organizations rather than by individual farmers to insure full participation in the program [ 46 ].
Pheromone-based control tactics including mass capturing of using pheromone traps Figure 8 proved to be effective against a variety of insect pests in area-wide IPM programs. Pheromone-baited trap for monitoring and mass trapping of red palm weevil. Successful IPM depends mainly on basic research on ecosystem and the understanding of interactions among hosts, pests, and their natural enemies [ 11 ].
The following steps should be taken before implementing an IPM program: identify the pest;. The socioeconomic factor is important in the implementation of IPM. For example, the decision to include a new variety resistant to insects may also depend on the market value of that variety. Preparation of guidelines that include the principles of IPM for different crops is essential during the implementation phase.
Moreover, the continuous evaluation of IPM programs provides feedback for future adjustment and improvement [ 27 ]. It is extremely important to record and evaluate the results of your control efforts. Some control methods, especially nonchemical procedures, are slow to yield measurable results.
Other methods may be ineffective or even damaging to the target crop, animal, treated surface, or natural predators and parasites. Pesticide use by volume, pesticide use by treatment frequency index, reduction in use of more toxic pesticides, and environmental impact quotient have been used as IPM impact evaluation indicators [ 22 ].
Since , no major departures from the basic notion of IPM have occurred [ 11 ]. In the future, major advances in IPM are expected in decision-making techniques as well as tactical options for control methods. New generation of GPS, sensors-fitted farm equipment, e-tablets, and mobile applications Plantix could be used for future pests and diseases identification and monitoring [ 47 ].
Since implementation of IPM programs depends largely on information, it is anticipated that a giant step being taken in areas such as principles of insect sampling, computer programming and mathematics, understanding of pests biological and ecological aspects, and simulation techniques and modeling [ 11 ].
Additionally, meteorological and geostatistical computer models can revolutionize forecasting and monitoring of insect pests, which, eventually improve decision-making for IPM. Novel tactics such as silencing of pest gene or RNA interference RNAi and endosymbionts hosted by insect pests could be used as potential new tools for future management of insect pests. Continuous training and education of farmers represent the cornerstone for establishment of solid and effective IPM program in agroecosystems.
Due to its importance, the European Union has adopted IPM as a policy for management of insects and other pests. Manipulating reproduction of insect pests with pheromones, irradiations, Wolbachia , and pathogens will provide a variety of innovative tactical methods for IPM.
Transgenic plants resistant to insect pest are also important tactical methods for future implementation of IPM. The information and communication technology ICT and nonprint media such as projectors, tablets, laptops, and mobile cell phones are expected to play a vital role in disseminating IPM knowledge among illiterate farmers, in their languages, in developing countries.
The advancement in semiochemical-based tactics could provide great support for area-wide IPM AW-IPM , which will gain importance in the coming years due to the increasing numbers of invasive insect pest species.
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Edited by Dalila Haouas. Edited by Farzana Khan Perveen. We are IntechOpen, the world's leading publisher of Open Access books. Built by scientists, for scientists. Our readership spans scientists, professors, researchers, librarians, and students, as well as business professionals. Downloaded: Abstract Insect pests cause substantial losses to food and fiber crops worldwide. Keywords insect pest integrated pest management economic threshold economic injury level decision rules ecosystem.
Introduction Insects appeared on earth about million years and have diversified into several million species that have adapted to almost all available ecosystems. Table 1. Because parasitoids must be adapted to the life cycle, physiology and defenses of their hosts, they are limited in their host range, and many are highly specialized. Thus, accurate identification of the host and parasitoid species is critically important in using parasitoids for biological control.
Pathogens : Insects, like other animals and plants, are infected by bacteria, fungi, protozoans and viruses that cause disease. These diseases may reduce the rate of feeding and growth of insect pests, slow or prevent their reproduction, or kill them. In addition, insects are also attacked by some species of nematodes that, with their bacterial symbionts, cause disease or death.
Under certain environmental conditions, diseases can multiply and spread naturally through an insect population, particularly when the density of the insects is high. An example of an established population of an insect pathogen which has been successfully controlling its host is the fungus Entomophaga maimaiga , a pathogen of the gypsy moth.
This fungus is believed to have been introduced about , but was not discovered in forests until , when it was widespread and abundant in New England. It has continued to control gypsy moth populations here for several years. It overwinters in leaf litter as resting spores, which germinate when gypsy moth larvae are present. First-instar caterpillars are dispersed by wind, and those that fall to the forest floor are probably infected while crawling to a tree.
While these larvae are feeding in the tree canopy, if there is adequate rainfall, the fungus in their bodies produces spores that spread to other caterpillars. If conditions are suitable, this infection cycle will occur again during the larval stage. Large caterpillars rest during the day in forest litter, where they are also susceptible to infection by germinating resting spores.
In late June, as infected caterpillars die in large numbers, new resting spores are produced to survive the next winter. This biological control agent is dependent on rain at appropriate times during the season to be successful. There are three primary methods of using biological control in the field: 1 conservation of existing natural enemies, 2 introducing new natural enemies and establishing a permanent population called "classical biological control" , and 3 mass rearing and periodic release, either on a seasonal basis or inundatively.
Reducing pesticide use: Most natural enemies are highly susceptible to pesticides, and pesticide use is a major limitation to their effectiveness in the field. The original idea that inspired integrated pest management IPM was to combine biological and chemical control by reducing pesticide use to the minimum required for economic production, and applying the required pesticides in a manner that is least disruptive to biological control agents. The need for pesticides can be reduced by use of resistant varieties, cultural methods that reduce pest abundance or damage, methods of manipulating pest mating or host-finding behavior, and, in some cases, physical methods of control.
Many IPM programs, however, have not been able to move beyond the first stage of developing sampling methods and economic thresholds for pesticide application. Although there are variations by crop and class of pesticide, the overall trend is that previous reductions, due to the substitution of economic thresholds for calendar spraying and the use of pesticides effective at lower dosages, are being reversed by increases in acreage treated and number of treatments per season.
This stagnation of pest management has resulted in calls for IPM to be re-focused toward preventing pest problems by greater understanding of pest ecology, enhancing the ability of plants and animals to defend themselves against pests, and building populations of beneficial organisms. This strategy is sometimes called "biointensive IPM. The effect of a pesticide on natural enemy populations depends on the physiological effect of the chemical and on how the pesticide is used -- how and when it is applied, for example.
While insecticides and acaricides are most likely to be toxic to insect and mite natural enemies, herbicides and fungicides are sometimes toxic as well.
A database has been compiled on the effects of pesticides on beneficial insects, spiders and mites summarized in Croft and Benbrook Among the insecticides, synthetic pyrethroids are among the most toxic to beneficials, while Bacillus thuringiensis and insect growth regulators were among the least toxic. In general, systemic insecticides, which require consuming plant material for exposure, and insecticides that must be ingested for toxicity affect natural enemies much less than pests.
Pesticides may also have more subtle effects on the physiology of natural enemies than direct toxicity. Several fungicides, such as benomyl, thiophanate-methyl, and carbendazim, inhibit oviposition by predacious phytoseiid mites. Certain herbicides diquat and paraquat make the treated soil in vineyards repellent to predacious mites.
The impact of pesticides on natural enemies can be reduced by careful timing and placement of applications to minimize contact between the beneficial organism and the pesticide. Less persistent pesticides reduce contact, especially if used with knowledge of the biology of the natural enemy to avoid susceptible life stages.
Spot applications in the areas of high pest density or treatment of alternating strips within a field may leave natural enemies in adjacent areas unaffected. The effectivenss of limiting the areas treated may depend on the mobility of the natural enemy and the pest. Natural enemies are generally not active during the winter in the Northeast, and thus, unless they are re-released each year, must have a suitable environment for overwintering.
Some parasitoids and pathogens overwinter in the bodies of their hosts which may then have overwintering requirements of their own , but others may pass the winter in crop residues, other vegetation, or in soil.
A classic example is the overwintering of predacious mites in fruit orchards. Ground cover in these orchards provides shelter over the winter, refuge from pesticides used on the fruit trees, and a source of pollen and alternate prey. The adults of many predators and parasitoids may require or benefit from pollen, nectar or honeydew produced by aphids during the summer. Many crop plants flower uniformly for only a short time, so flowering plants along the edges of the field or within the field may be needed as supplemental sources of pollen and nectar.
However, diversification of plants within the field can also interfere with the efficiency of host-finding, particularly for specialist parasitoids. Populations of generalist predators may be stabilized by the availability of pollen and alternative prey, but the effectiveness of the predators still depends on whether they respond quickly enough, either by aggregation or multiplication, to outbreaks of the target pest.
Thus, diversification of plants or other methods of supplementing the nutrition of natural enemies must be done with knowledge of the behavior and biology of the natural enemy and pest. For example, the native lady beetle Coleomegilla maculata is a potentially important predator of the eggs and early instar larvae of Colorado potato beetle. The population feeding on the potato beetle depends on the availability of aphid prey in surrounding fields, including crops of alfalfa, brassicas, cucurbits, and corn, and on the availability of pollen from corn and several weeds, such as dandelion and yellow rocket.
Although this predator does not currently control Colorado potato beetle on its own, more knowledge about managing C. This is a process which requires extensive research into the biology of the pest, potential natural enemies and their biology, and the possibility of unintended consequences e.
After suitable natural enemies are found, studied, and collected, they must undergo quarantine to eliminate any pathogens or parasites on the natural enemy population. Then, the natural enemies are carefully released, with attention to proper timing in the enemy and pest life cycles, in a site where the target pest is abundant, and where disturbance of the newly released enemies is minimized. Although this process is long and complex, when it is successful, the results can be impressive and permanent, as long as care is taken in production practices to minimize negative effects on the natural enemy.
One of many examples of a pest controlled by successful introduction of new natural enemies is the alfalfa weevil. The alfalfa weevil is native to Europe, and was first reported in the US in Larval densities were high enough to require most growers to spray one or more times per year. Several parasitoids were introduced from Europe against this pest.
The most successful introductions include two species of parasitoids attacking the larvae, one attacking the adult, and a parasitoid and predator attacking the eggs.
Schematic showing design of the population suppression and insecticide resistance management experiments. The glasshouse cage population suppression experiment was designed to simulate a field scenario in which biotic and abiotic mortality factors maintain a stable pest population, with adult moths mating and laying eggs in glasshouse cages, and larval rearing being conducted in a separate temperature-controlled room [ 18 , 20 , 27 ].
When weekly OXL male moth releases started, OXL-treated cages were paired with an untreated counterpart control cage and the number of pupae subsequently reintroduced into each treatment cage was made proportional to the ratio of eggs counted between the treatment cage and its paired control cage: this allowed the expected reduced number of females in OXL-treated cages to be reflected in reduced numbers of pupae entering the population in the next generation.
In contrast, the subsequent glasshouse cage IRM experiment was conducted on broccoli plants — some on Bt broccoli — as they offered a realistic model for many agricultural systems with transgenic crops. Artificially maintaining stable populations on plants, by accurately manipulating the number of progeny surviving, was deemed impractical, and the populations were therefore allowed to expand freely in the near-absence of biotic and abiotic mortality factors, with all life stages of each experimental population residing in their respective cage.
This second experiment provided a more rigorous test of the pest management potential of the OXL strain as the intrinsic growth rate of target populations was not artificially controlled and competitors from these target populations had the advantage of being reared on host plant material, as opposed to an artificial diet medium.
Generations in the population suppression experiment were continuous while those in the resistance management experiment were discrete, at least within the experimental period.
In the population suppression experiment a continuous generational structure was achieved by introducing pupae into cages three times per week. As the lifecycle of P. In the resistance management experiment conducted on broccoli, populations were founded by a single introduction of adult moths. Adults in this founding generation, and those in subsequent generations, were allowed to mate freely in their respective cages with females from the same generation ovipositing over a number of days.
As such, within the experimental period generations remained discrete and were discernable by peaks in weekly population counts which, as expected, reached a maximum level once per generation approximately every 3 weeks; see below. Generations did, however, begin to overlap over time. The generation time of P. As experiments were conducted in a glasshouse, these time periods varied depending on outdoor weather conditions. Experiments were conducted in quarantine facilities at Rothamsted Research, Hertfordshire, UK in accordance with legislation concerning the contained use of GM organisms in the UK.
As an efficient, easily replicable proxy for host crop infestation, larval rearing was conducted on artificial diet medium and followed the methods of Martins et al. All experimental populations were reared on non-tetracycline diet. Insects were added to the cages as pupae. The experiment comprised two phases: establishment and suppression. During establishment, stable mixed-age populations of wild-type P.
During suppression, weekly introductions of transgene-homozygous OXL males were made into two of the cages to investigate whether engineered female-specific lethality resulted in suppression of these populations. Throughout the experiment, cabbage-extract-baited Parafilm Bemis Company Inc. These were replaced three times per week Monday, Wednesday, and Friday and eggs collected on each sheet were counted. During each egg collection, dead adult moths and uneclosed pupae were also collected from the cages, sexed, and counted.
Eclosed adults were provided with sugar water-saturated cotton wool reservoirs, changed every 2—3 days. Cage populations were initiated by placing unsexed wild-type pupae into each cage. Stable non-expanding populations were maintained in each cage to mimic the stabilizing effects of predation and other limiting factors in the wild.
This was achieved by introducing a constant number of pupae back into the cages each week. A total of first-instar larvae were selected each week to carry on the population, taken from the three weekly collections 67 larvae chosen from Monday collection, 67 from Wednesday, and 66 from Friday.
These larvae were reared in plastic beakers on non-tetracycline diet, and after pupation were sexed and transferred back into the cage from which they had been collected as eggs, 2 weeks earlier. Tri-weekly egg collections and subsequent tri-weekly pupal reintroductions maintained stable, mixed-age populations as might be expected in the field, where adult moths would be continuously entering the population.
The first two introductions of wild-type moths into each cage weeks 1 and 2 originated from an independent laboratory colony. From this point onwards each cage population was self-sustaining. Once egg counts had stabilized indicating stable populations in week 9, two of the four cages were chosen as OXL treatment cages Cages 2 and 4 and two as control cages Cages 1 and 3. Cages were designated in a blocked design to minimize bias caused by uncontrolled abiotic factors. In non-treatment control cages, the protocol from the establishment phase was continued.
Each OXL treatment cage was randomly paired with a control cage for the remainder of the experiment Cages 1 and 2; Cages 3 and 4. The reintroduction of pupae into each OXL treatment cage followed the same protocol as for the establishment phase; however, the total number of early-instar larvae selected for rearing that week was made proportional to the ratio between the number of eggs collected for that treatment cage and its paired control cage for the week when these larvae were collected as eggs.
This method ensured that the population suppression effect of female death of transgenic larvae, reflected later by reduced number of eggs collected, resulted in reduced numbers of pupae re-entering the OXL-treated cages relative to an untreated population. After cages had been paired, weekly introductions of homozygous OXL males into the treatment cages began. The target over-flooding ratio OXL to wild-type males entering the population was set at This release rate is in line with previous studies investigating the effect of female-lethal transgenes on caged populations [ 18 , 20 , 27 ] and much lower than the sterile:wild ratios that successful SIT programs have aimed to achieve against other Lepidoptera: P.
Males were released as pupae into the cages once per week Wednesday. Pupae reintroduced into treatment cages were screened for the DsRed2 fluorescent protein transformation marker and fluorescence proportions proportion of the population which carried the transgene recorded. Under the restrictive conditions of this experiment and the highly penetrant female-lethal phenotype of OXL [ 15 ], population-level transgene allele frequencies were equal to half the fluorescence proportion [ 48 ].
All P. Eleven cages, each 1. Treatments 1—3 were assigned three cages replicates while Treatment 4 was assigned two cages. A total of males and females from this cross were then mated to produce the founder strain. These crosses provide an expected resistance allele frequency of 0. However, Bt survival assay results Fig.
Two strains of broccoli Brassica oleracea L. Together, this transgenic plant cultivar and the Cry1-Ac-resistant P. Cry1-Ac toxin production was verified by screening the plants 4—5 weeks old with susceptible Geneva 88 strain neonates. All cages started with 20 broccoli plants of their respective cultivar. Plants were replaced after 4 weeks, or when defoliated due to larval feeding, by cutting them at their base and placing them on the replacement plants to allow larvae to transfer.
In all treatments, replicates were initiated by the release of P. In treatments involving Bt plants, randomly selected adults were released. In treatments involving non- Bt plants, seven males and females total 14 adults were released into the cages.
Due to Bt selection, this gave approximately equal starting population densities in Generation 1 in all cages. OXL-homozygous males for release into the cages were produced by rearing egg collections from a stock colony in the absence of tetracycline in larval feed. Releases in this experiment were proportional: daily estimates of adult male recruitment for each cage were used to calculate a daily male release number for that cage, dependent on the release rate pre-determined for that treatment.
A proportional release rate, rather than a constant number, was applied to reduce the likelihood of population extinction and thereby allow exploration of the effect of MS transgene releases on population dynamics and resistance allele frequency. Release rates were selected in advance based on the outcomes of predictive deterministic models, investigating the effects of each experimental treatment on population size. In Treatments 2 and 3 low-OXL release treatments , the intended over-flooding ratio was transgenic: wild-type males.
However, due to insect rearing limitations this was limited to in Generation 1, and increased to thereafter. These low release rates, in combination with Bt plants, were predicted to maintain a relatively constant population size, but insufficient to cause population suppression when used alone. Similarly, the over-flooding rates in cages with high-OXL releases, initially and increased to in Generation 2 as production capacity increased, were predicted to be sufficient for population suppression when used alone.
Where small populations are expanding in the absence of limiting biotic or abiotic factors, stochastic effects make it difficult to accurately predict the rate of population increase. In response to higher-than-predicted population growth and limited plants available, the cages with Bt broccoli plants and low-rate weekly OXL releases were reduced to two replicates in Generation 3 terminated cage selected at random. In Generation 4, the number of plants in each treatment with Bt broccoli only no OXL releases cage was reduced to five while maintaining per-plant population density levels in response to limited plant availability.
As generations were still discrete, this manipulation was timed during a period when the vast majority of insects in the population were present as larvae or pupae on plants, allowing accurate culling of the population.
In Generation 3, requirements for OXL male moths exceeded production capacity, so the treatment with high-rate weekly OXL releases was reduced to one replicate only. The numbers of second to fourth instar larvae and pupae on each plant in each cage were counted once per week.
With the exception of cages with high-rate OXL releases, Bt resistance data were collected from all cages in their final generation. As each cage reached maximum egg-laying potential judged by female recruitment data in each population, collected from eclosion cages , eggs were collected from 8—10 leaves selected at random from each cage. For each cage, two replicated Bt survival assays and one control assay were performed. Non- Bt controls were prepared in the same way, with no added Bt solution.
At third instar, larvae were transferred onto the air-dried diet surface. Exact numbers of larvae per replicate differed between cages according to availability of eggs. A minimum of 33 larvae per pot were used for the first Bt replicate; 11 per pot for the second Bt replicate; and all control replicates contained 20 larvae. Mortality was assessed 72 h later with surviving larvae taken to be homozygous for the resistance allele. For comparison, the founder strain was subjected to the same assay, in the generation prior to experimental initiation.
This reduction in survival was used to calibrate the results of the assay for other treatments prior to calculation of allele frequencies.
Once calibrated, the square of the proportion surviving gave an estimate of the Bt allele frequency in the founder strain under Hardy-Weinberg equilibrium. The validity of this assumption was explored by comparing the percentage of non-surviving individuals with DsRed2 fluorescence proportions. The population-level fluorescence proportion proportion of the population carrying the transgene was estimated for all cages into which OXL males were released, in the final generations they were conducted.
The exception to this was the high OXL release treatment where this assay was conducted in the penultimate third generation due to the small population size of this treatment cage in Generation 4. Each cage acted as a replicate with a minimum of 75 larvae per assay pot. These were placed into smaller cages in an adjacent glasshouse maintained in the same environmental conditions. The number and sex ratio of the adults eclosing from these plants were recorded daily, and eclosed adults were then returned to their respective experimental cages.
From this data an estimate of the eclosion rates in the main cage was made, from which the number of OXL males required to achieve the respective over-flooding ratio for each cage was calculated. For Generation 4, a t -test was used as only two treatments were compared. For Bt assay results, results from the two Bt assays within each cage were summed to avoid pseudo-replication and then corrected for control mortality using a Henderson-Tilton correction. For comparisons between treatments, a Logit model for Categorical Data Analysis was used followed by pair-wise comparisons using Generalized Linear Hypothesis Testing with each cage acting as a replicate.
Data were analyzed using R Version 2. Lounibos LP. Invasions by insect vectors of human disease. Annu Rev Entomol. Ecological effects of invasive alien insects. J Biol Inv. Article Google Scholar. Oerke E-C. Crop losses to pests. J Agric Sci. All Reviews:. Popular user-defined tags for this product:.
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