Insecticide

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FLIT manual spray pump from 1928 FLIT Spray Can 1.jpg
FLIT manual spray pump from 1928
Farmer spraying a cashewnut tree in Tanzania Kente l.jpg
Farmer spraying a cashewnut tree in Tanzania

Insecticides are pesticides used to kill insects. [1] They include ovicides and larvicides used against insect eggs and larvae, respectively. The major use of insecticides is in agriculture, but they are also used in home and garden settings, industrial buildings, for vector control, and control of insect parasites of animals and humans.

Contents

Acaricides, which kill mites and ticks, are not strictly insecticides, but are usually classified together with insecticides. Some insecticides (including common bug sprays) are effective against other non-insect arthropods as well, such as scorpions, spiders, etc. Insecticides are distinct from insect repellents, which repel but do not kill.

Sales

In 2016 insecticides were estimated to account for 18% of worldwide pesticide sales. [2] Worldwide sales of insecticides in 2018 were estimated as $ 18.4 billion, of which 25% were neonicotinoids, 17% were pyrethroids, 13% were diamides, and the rest were many other classes which sold for less than 10% each of the market. [3]

Synthetic insecticides

Insecticides are most usefully categorised according to their modes of action. The insecticide resistance action committee (IRAC) list s 30 modes of action plus unknowns. There can be several chemical classes of insecticide with the same mode or action. IRAC lists 56 chemical classed plus unknowns. Further Information: List of insecticides.

The mode of action describes how the insecticide kills or inactivates a pest. It provides another way of classifying insecticides.

Development

Insecticides with systemic activity against sucking pests, which are safe to pollinators, are sought after, [4] [5] [6] particularly in view of the partial bans on neonicotinoids. Revised 2023 guidance by registration authorities describes the bee testing that is required for new insecticides to be approved for commercial use. [7] [8] [9] [10]

Systemicity and Translocation

Insecticides may be systemic or non-systemic (contact insecticides). [2] [11] [12] Systemic insecticides penetrate into the plant and move (translocate) inside the plant. Translocation may be upward in the xylem, or downward in the phloem or both. Systemicity is a prerequisite for the pesticide to be used as a seed-treatment. Contact insecticides (non-systemic insecticides) remain on the leaf surface and act through direct contact with the insect.

Insects feed from various compartments in the plant. Most of the major pests are either chewing insects or sucking insects. [13] Chewing insects, such as caterpillars, eat whole pieces of leaf. Sucking insects use feeding tubes to feed from phloem (e.g. aphids, leafhoppers, scales and whiteflies), or to suck cell contents (e.g. thrips and mites). An insecticide is more effective if it is in the compartment the insect feeds from. The physicochemical properties of the insecticide determine how it is distributed throughout the plant. [11] [12]

Organochlorides

The best known organochloride, DDT, was created by Swiss scientist Paul Müller. For this discovery, he was awarded the 1948 Nobel Prize for Physiology or Medicine. [14] DDT was introduced in 1944. It functions by opening sodium channels in the insect's nerve cells. [15] The contemporaneous rise of the chemical industry facilitated large-scale production of chlorinated hydrocarbons including various cyclodiene and hexachlorocyclohexane compounds. Although commonly used in the past, many older chemicals have been removed from the market due to their health and environmental effects (e.g. DDT, chlordane, and toxaphene). [16] [17]

Organophosphates

Organophosphates are another large class of contact insecticides. These also target the insect's nervous system. Organophosphates interfere with the enzymes acetylcholinesterase and other cholinesterases, causing an increase in synaptic acetylcholine and overstimulation of the parasympathetic nervous system. [18] and killing or disabling the insect. Organophosphate insecticides and chemical warfare nerve agents (such as sarin, tabun, soman, and VX) have the same mechanism of action. Organophosphates have a cumulative toxic effect to wildlife, so multiple exposures to the chemicals amplifies the toxicity. [19] In the US, organophosphate use declined with the rise of substitutes. [20] Many of these insecticides, first developed in the mid 20th century, are very poisonous. [21] Many organophosphates do not persist in the environment.

Carbamates

Carbamate insecticides have similar mechanisms to organophosphates, but have a much shorter duration of action and are somewhat less toxic.[ citation needed ]

Pyrethroids

Pyrethroid insecticides mimic the insecticidal activity of the natural compound pyrethrin, the biopesticide found in Pyrethrum (Now Chrysanthemum and Tanacetum ) species. They have been modified to increase their stability in the environment. These compounds are nonpersistent sodium channel modulators and are less toxic than organophosphates and carbamates. Compounds in this group are often applied against household pests. [22] Some synthetic pyrethroids are toxic to the nervous system. [23]

Neonicotinoids

Neonicotinoids are a class of neuro-active insecticides chemically similar to nicotine.(with much lower acute mammalian toxicity and greater field persistence). These chemicals are acetylcholine receptor agonists. They are broad-spectrum systemic insecticides, with rapid action (minutes-hours). They are applied as sprays, drenches, seed and soil treatments. Treated insects exhibit leg tremors, rapid wing motion, stylet withdrawal (aphids), disoriented movement, paralysis and death. [24] Imidacloprid, of the neonicotinoid family, is the most widely used insecticide in the world. [25] In the late 1990s neonicotinoids came under increasing scrutiny over their environmental impact and were linked in a range of studies to adverse ecological effects, including honey-bee colony collapse disorder (CCD) and loss of birds due to a reduction in insect populations. In 2013, the European Union and a few non EU countries restricted the use of certain neonicotinoids. [26] [27] [28] [29] [30] [31] [32] [33] and its potential to increase the susceptibility of rice to planthopper attacks. [34]

Phenylpyrazoles

Phenylpyrazole insecticides, such as fipronil are a class of synthetic insecticides that operate by interfering with GABA receptors. [35]

Butenolides

Butenolide pesticides are a novel group of chemicals, similar to neonicotinoids in their mode of action, that have so far only one representative: flupyradifurone. They are acetylcholine receptor agonists, like neonicotinoids, but with a different pharmacophore. [36] They are broad-spectrum systemic insecticides, applied as sprays, drenches, seed and soil treatments. Although the classic risk assessment considered this insecticide group (and flupyradifurone specifically) safe for bees, novel research [37] has raised concern on their lethal and sublethal effects, alone or in combination with other chemicals or environmental factors. [38] [39]

Diamides

Diamides selectively activate insect ryanodine receptors (RyR), which are large calcium release channels present in cardiac and skeletal muscle, [40] leading to the loss of calcium crucial for biological processes. This causes insects to act lethargic, stop feeding, and eventually die. [41] The first insecticide from this class to be registered was flubendiamide. [41]

Insect growth regulators

Insect growth regulator (IGR) is a term coined to include insect hormone mimics and an earlier class of chemicals, the benzoylphenyl ureas, which inhibit chitin (exoskeleton) biosynthesis in insects [42] Diflubenzuron is a member of the latter class, used primarily to control caterpillars that are pests. Of these, methoprene is most widely used. It has no observable acute toxicity in rats and is approved by World Health Organization (WHO) for use in drinking water cisterns to combat malaria. Most of its uses are to combat insects where the adult is the pest, including mosquitoes, several fly species, and fleas. Two very similar products, hydroprene and kinoprene, are used for controlling species such as cockroaches and white flies. Methoprene was registered with the EPA in 1975. Virtually no reports of resistance have been filed. A more recent type of IGR is the ecdysone agonist tebufenozide (MIMIC), which is used in forestry and other applications for control of caterpillars, which are far more sensitive to its hormonal effects than other insect orders.

Biological pesticides

Definition

The EU defines biopesticides as "a form of pesticide based on micro-organisms or natural products". [43] The US EPA defines biopesticides as “certain types of pesticides derived from such natural materials as animals, plants, bacteria, and certain minerals”. [44] Microorganisms that control pests may also be categorised as biological pest control agents together with larger organisms such as parasitic insects, entomopathic nematodes etc. Natural products may also be categorised as chemical insecticides.

The US EPA describes three types of biopesticide. [44] Biochemical pesticides (meaning bio-derived chemicals), which are naturally occurring substances that control pests by non-toxic mechanisms. Microbial pesticides consisting of a microorganism (e.g., a bacterium, fungus, virus or protozoan) as the active ingredient. Plant-Incorporated-Protectants (PIPs) are pesticidal substances that plants produce from genetic material that has been added to the plant (thus producing transgenic crops).

Market

The global bio-insecticide market was estimated to be less than 10% of the total insecticide market. [45] The bio-insecticde market is dominated by microbials. [46] The bio-insecticide market is growing more that 10% yearly, which is a higher growth than the total insecticide market, mainly due to the increase in organic farming and IPM, and also due to benevolent government policies. [45]

Biopesticides are regarded by the US and European authorities as posing fewer risks of environmental and mammalian toxicity. [44] Biopesticides are more than 10 x (often 100 x) cheaper and 3 x faster to register than synthetic pesticides. [45]

Advantages and disadvantages

There is a wide variety of biological insecticides with differing attributes, but in general the following has been described. [47] [48]

They are easier, faster and cheaper to register, usually with lower mammalian toxicity. They are more specific, and thus preserve beneficial insects and biodiversity in general. This makes them compatible with IPM regimes. They degrade rapidly cause less impact on the environment. They have a shorter withholding period.

The spectrum of control is narrow. They are less effective and prone to adverse ambient conditions. They degrade rapidly and are thus less persistant. They are slower to act. They are more expensive, have a shorter shelf-life, and are more difficult to source. They require mor specialised knowledge to use.

Plant Extracts

Most or all plants produce chemical insecticides to stop insects eating them. Extracts and purified chemicals from thousands of plants have been shown to be insecticidal, however only a few are used in agriculture. [49] In the USA 13 are registered for use, in the EU 6. In Korea, where it is easier to register botanical pesticides, 38 are used. Most used are neem oil, chenopodium, pyrethrins, and azadirachtin. [49] Many botanical insecticides used in past decades (e.g. rotenone, nicotine, ryanodine) have been banned because of their toxicity. [49]

Genetically modified crops

The first transgenic crop, which incorporated an insecticidal PIP, contained a gene for the CRY toxin from Bacillus thuringiensis (B.t.) and was introduced in 1997. [50] For the next ca 25 years the only insecticidal agents used in GMOs were the CRY and VIP toxins from various strains of B.t, which control a wide number of insect types. These are widely used with > 100 million hectares planted with B.t. modified crops in 2019. [50] Since 2020 several novel agents have been engineered into plants and approved.  ipd072Aa from Pseudomonas chlororaphis, ipd079Ea from Ophioglossum pendulum, and mpp75Aa1.1 from Brevibacillus laterosporus code for protein toxins. [50] [51] The trait dvsnf7 is an RNAi agent consisting of a double-stranded RNA transcript containing a 240 bp fragment of the WCR Snf7 gene of the western corn rootworm (Diabrotica virgifera virgifera). [51] [52]

RNA interference

RNA interference (RNAi) uses segments of RNA to fatally silence crucial insect genes. [53] In 2024 two uses of RNAi have been registered by the authorities for use:Genetic modification of a crop to introduce a gene coding for an RNAi fragment, and spraying double stranded RNA fragments onto a field. [52] Monsanto introduced the trait DvSnf7 which expresses a double-stranded RNA transcript containing a 240 bp fragment of the WCR Snf7 gene of the Western Corn Rootworm. [51] GreenLight Biosciences introduced Ledprona, a formulation of double stranded RNA as a spray for potato fields. It targets the essential gene for proteasome subunit beta type-5 (PSMB5) in the Colorado potato beetle. [52]

Spider toxins

Spider venoms contain many, often hundreds, of insecticidally active toxins. Many are proteins that attack the nervous system of the insect. [54] Vestaron introduced for agricultural use a spray formulation of GS-omega/kappa-Hxtx-Hv1a (HXTX), derived from the venom of the Australian blue mountain funnel web spider (Hadronyche versuta). [54]

Entomopathic bacteria

Entomopathic bacteria can be mass-produced. [46] The most widely used is Bacillus thuringiensis (B.t.), used since decades. There are several strains used with different applications against lepidoptera, coleoptera and diptera. Also used are Lysinibacillus sphaericus, Burkholderia spp, and Wolbachia pipientis. Avermectins and spinosyns are bacterial metabolites, mass-produced by fermentation and used as insecticides. The toxins from B.t. have been incorporated into plants through genetic engineering. [46]

Entomopathic fungi

Entomopathic fungi have been used since 1965 for agricultural use. Hundreds of strains are now in use. They often kill a broad range of insect species. Most strains are from Beauveria, Metarhizium, Cordyceps and Akanthomyces species. [55]

Entomopathic viruses

Of the many types of entomopathic viruses, only baculaviruses are used commercially, and are each specific for their target insect. They have to be grown on insects, so their production is labour-intensive. [56]

Environmental toxicity

Effects on nontarget species

Some insecticides kill or harm other creatures in addition to those they are intended to kill. For example, birds may be poisoned when they eat food that was recently sprayed with insecticides or when they mistake an insecticide granule on the ground for food and eat it. [19] Sprayed insecticide may drift from the area to which it is applied and into wildlife areas, especially when it is sprayed aerially. [19]

Persistence in the environment and accumulation in the food chain

DDT was the first organic insecticide. It was introduced during WW2, and was widely used. One use was vector control and it was sprayed on open water. It degrades slowly in the environment, and it is lipophilic (fat soluble). It became the first global pollutant, and the first pollutant to accumulate [57] and magnify in the food chain. [58] [59] During the 1950s and 1960s these very undesirable side effects were recognized, and after some often contentious discussion, DDT was banned in many countries in the 1960s and 1970s. Finally in 2001 DDT and all other persistent insecticides were banned via the Stockholm Convention. [60] [61] Since many decades the authorities require new insecticides to degrade in the environment and not to bioaccumulate. [62]

Runoff and percolation

Solid bait and liquid insecticides, especially if improperly applied in a location, get moved by water flow. Often, this happens through nonpoint sources where runoff carries insecticides in to larger bodies of water. As snow melts and rainfall moves over and through the ground, the water picks applied insecticides and deposits them in to larger bodies of water, rivers, wetlands, underground sources of previously potable water, and percolates in to watersheds. [63] This runoff and percolation of insecticides can effect the quality of water sources, harming the natural ecology and thus, indirectly effect human populations through biomagnification and bioaccumulation.

Insect decline

Both number of insects and number of insect species have declined dramatically and continuously over past decades, causing much concern. [64] [65] [66] Many causes are proposed to contribute to this decline, the most agreed upon are loss of habitat, intensification of farming practices, and insecticide usage. Domestic bees were declining some years ago [67] but population and number of colonies have now risen both in the USA [68] and worldwide. [69] Wild species of bees are still declining.

Bird decline

Besides the effects of direct consumption of insecticides, populations of insectivorous birds decline due to the collapse of their prey populations. Spraying of especially wheat and corn in Europe is believed to have caused an 80 per cent decline in flying insects, which in turn has reduced local bird populations by one to two thirds. [70]

Alternatives

Instead of using chemical insecticides to avoid crop damage caused by insects, there are many alternative options available now that can protect farmers from major economic losses. [71] Some of them are:

  1. Breeding crops resistant, or at least less susceptible, to pest attacks. [72]
  2. Releasing predators, parasitoids, or pathogens to control pest populations as a form of biological control. [73]
  3. Chemical control like releasing pheromones into the field to confuse the insects into not being able to find mates and reproduce. [74]
  4. Integrated Pest Management: using multiple techniques in tandem to achieve optimal results. [75]
  5. Push-pull technique: intercropping with a "push" crop that repels the pest, and planting a "pull" crop on the boundary that attracts and traps it. [76]

Examples

Source: [77]

Organochlorides

Organophosphates

Carbamates

Pyrethroids

Neonicotinoids

Ryanoids

Insect growth regulators

Derived from plants or microbes

Biologicals

Inorganic/mineral derived insecticides

See also

Related Research Articles

<i>Bacillus thuringiensis</i> Species of bacteria used as an insecticide

Bacillus thuringiensis is a gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. B. thuringiensis also occurs naturally in the gut of caterpillars of various types of moths and butterflies, as well on leaf surfaces, aquatic environments, animal feces, insect-rich environments, flour mills and grain-storage facilities. It has also been observed to parasitize moths such as Cadra calidella—in laboratory experiments working with C. calidella, many of the moths were diseased due to this parasite.

<span class="mw-page-title-main">Pesticide</span> Substance used to destroy pests

Pesticides are substances that are used to control pests. They include herbicides, insecticides, nematicides, fungicides, and many others. The most common of these are herbicides, which account for approximately 50% of all pesticide use globally. Most pesticides are used as plant protection products, which in general protect plants from weeds, fungi, or insects. In general, a pesticide is a chemical or biological agent that deters, incapacitates, kills, or otherwise discourages pests. Target pests can include insects, plant pathogens, weeds, molluscs, birds, mammals, fish, nematodes (roundworms), and microbes that destroy property, cause nuisance, or spread disease, or are disease vectors. Along with these benefits, pesticides also have drawbacks, such as potential toxicity to humans and other species.

<span class="mw-page-title-main">Pesticide resistance</span> Decreased effectiveness of a pesticide on a pest

Pesticide resistance describes the decreased susceptibility of a pest population to a pesticide that was previously effective at controlling the pest. Pest species evolve pesticide resistance via natural selection: the most resistant specimens survive and pass on their acquired heritable changes traits to their offspring. If a pest has resistance then that will reduce the pesticide's efficacy – efficacy and resistance are inversely related.

<span class="mw-page-title-main">Pyrethrin</span> Class of organic chemical compounds with insecticidal properties

The pyrethrins are a class of organic compounds normally derived from Chrysanthemum cinerariifolium that have potent insecticidal activity by targeting the nervous systems of insects. Pyrethrin naturally occurs in chrysanthemum flowers and is often considered an organic insecticide when it is not combined with piperonyl butoxide or other synthetic adjuvants. Their insecticidal and insect-repellent properties have been known and used for thousands of years.

<span class="mw-page-title-main">Pyrethroid</span> Class of insecticides

A pyrethroid is an organic compound similar to the natural pyrethrins, which are produced by the flowers of pyrethrums. Pyrethroids are used as commercial and household insecticides.

<span class="mw-page-title-main">Imidacloprid</span> Chemical compound

Imidacloprid is a systemic insecticide belonging to a class of chemicals called the neonicotinoids which act on the central nervous system of insects. The chemical works by interfering with the transmission of stimuli in the insect nervous system. Specifically, it causes a blockage of the nicotinergic neuronal pathway. By blocking nicotinic acetylcholine receptors, imidacloprid prevents acetylcholine from transmitting impulses between nerves, resulting in the insect's paralysis and eventual death. It is effective on contact and via stomach action. Because imidacloprid binds much more strongly to insect neuron receptors than to mammal neuron receptors, this insecticide is more toxic to insects than to mammals.

<span class="mw-page-title-main">Fipronil</span> Chemical compound

Fipronil is a broad-spectrum insecticide that belongs to the phenylpyrazole chemical family. Fipronil disrupts the insect central nervous system by blocking the ligand-gated ion channel of the GABAA receptor and glutamate-gated chloride (GluCl) channels. This causes hyperexcitation of contaminated insects' nerves and muscles. Fipronil's specificity towards insects is believed to be due to its greater binding affinity for the GABAA receptors of insects than to those of mammals, and for its action on GluCl channels, which do not exist in mammals. As of 2017, there does not appear to be significant resistance among fleas to fipronil.

A Biopesticide is a biological substance or organism that damages, kills, or repels organisms seens as pests. Biological pest management intervention involves predatory, parasitic, or chemical relationships.

<span class="mw-page-title-main">Deltamethrin</span> Chemical compound

Deltamethrin is a pyrethroid ester insecticide. Deltamethrin plays a key role in controlling malaria vectors, and is used in the manufacture of long-lasting insecticidal mosquito nets; however, resistance of mosquitos and bed bugs to deltamethrin has seen a widespread increase.

Neonicotinoids are a class of neuro-active insecticides chemically similar to nicotine, developed by scientists at Shell and Bayer in the 1980s.

<span class="mw-page-title-main">Nitenpyram</span> Insecticide

Nitenpyram is a chemical frequently used as an insecticide in agriculture and veterinary medicine. The compound is an insect neurotoxin belonging to the class of neonicotinoids which works by blocking neural signaling of the central nervous system. It does so by binding irreversibly to the nicotinic acetylcholine receptor (nACHr) causing a stop of the flow of ions in the postsynaptic membrane of neurons leading to paralysis and death. Nitenpyram is highly selective towards the variation of the nACHr which insects possess, and has seen extensive use in targeted, insecticide applications.

<span class="mw-page-title-main">Environmental impact of pesticides</span> Environmental effect

The environmental effects of pesticides describe the broad series of consequences of using pesticides. The unintended consequences of pesticides is one of the main drivers of the negative impact of modern industrial agriculture on the environment. Pesticides, because they are toxic chemicals meant to kill pest species, can affect non-target species, such as plants, animals and humans. Over 98% of sprayed insecticides and 95% of herbicides reach a destination other than their target species, because they are sprayed or spread across entire agricultural fields. Other agrochemicals, such as fertilizers, can also have negative effects on the environment.

<span class="mw-page-title-main">Clothianidin</span> Chemical compound

Clothianidin is an insecticide developed by Takeda Chemical Industries and Bayer AG. Similar to thiamethoxam and imidacloprid, it is a neonicotinoid. Neonicotinoids are a class of insecticides that are chemically similar to nicotine, which has been used as a pesticide since the late 1700s. Clothianidin and other neonicotinoids act on the central nervous system of insects as an agonist of nAChR, the same receptor as acetylcholine, the neurotransmitter that stimulates and activating post-synaptic acetylcholine receptors but not inhibiting AChE. Clothianidin and other neonicotinoids were developed to last longer than nicotine, which is more toxic and which breaks down too quickly in the environment.

This is an index of articles relating to pesticides.

<span class="mw-page-title-main">Acetamiprid</span> Chemical compound

Acetamiprid is an organic compound with the chemical formula C10H11ClN4. It is an odorless neonicotinoid insecticide produced under the trade names Assail, and Chipco by Aventis CropSciences. It is systemic and intended to control sucking insects (Thysanoptera, Hemiptera, mainly aphids) on crops such as leafy vegetables, citrus fruits, pome fruits, grapes, cotton, cole crops, and ornamental plants. It is also a key pesticide in commercial cherry farming due to its effectiveness against the larvae of the cherry fruit fly.

<span class="mw-page-title-main">Naled</span> Organophosphate insecticide

Naled (Dibrom) is an organophosphate insecticide. Its chemical name is dimethyl 1,2-dibromo-2,2-dichloroethylphosphate.

<span class="mw-page-title-main">Thiamethoxam</span> Chemical compound

Thiamethoxam is the ISO common name for a mixture of cis-trans isomers used as a systemic insecticide of the neonicotinoid class. It has a broad spectrum of activity against many types of insects and can be used as a seed dressing.

<span class="mw-page-title-main">Fenpropathrin</span> Chemical compound

Fenpropathrin, or fenopropathrin, is a widely used pyrethroid insecticide in agriculture and household. Fenpropathrin is an ingestion and contact synthetic pyrethroid. Its mode of action is similar to other natural (pyrethrum) and synthetic pyrethroids where in they interfere with the kinetics of voltage gated sodium channels causing paralysis and death of the pest. Fenpropathrin was the first of the light-stable synthetic pyrethroids to be synthesized in 1971, but it was not commercialized until 1980. Like other pyrethroids with an α-cyano group, fenpropathrin also belongs to the termed type II pyrethroids. Type II pyrethroids are a more potent toxicant than type I in depolarizing insect nerves. Application rates of fenpropathrin in agriculture according to US environmental protection agency (EPA) varies by crop but is not to exceed 0.4 lb ai/acre.

<span class="mw-page-title-main">Flupyradifurone</span> Chemical compound

Flupyradifurone is a systemic butenolide insecticide developed by Bayer CropScience under the name Sivanto. Flupyradifurone protects crops from sap-feeding pests such as aphids and is safer for non-target organisms compared to other insecticides. Sivanto was launched in 2014 since it obtained its first commercial registration in central America. Insecticide Resistance Action Committee (IRAC) classified Flupyradifurone as 4D subset (butenolide) and it is the first pesticide in the butenolide category. It was approved by European Union in 2015.

Pesticide induced resurgence, often shortened to resurgence in pest management contexts, can be described as a constraint of pesticide use, by which they fail to control pests such as insects and spider mites: instead ‘flaring up’ populations that may have been of minor importance. Although there are more than one mechanisms by which this takes place, mortality of natural enemies following the use of broad-spectrum insecticides and acaricides is often implicated. This is sometimes called the “pesticide treadmill”: a term coined by Robert van den Bosch to describe a self-reinforcing over-dependence on agrochemicals and inimical to natural biological controls. Notable examples include the flare-up of rice brown planthopper populations, following over-use of broad-spectrum inscticides.

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