Aug 21, 2024
Understanding the Evolution of Resistance in Insect
A species' persistence requires the ability to adapt to the biotic and/or abiotic conditions present within its environment. This adaptation may occur through natural selection, which is the process by which organisms that are better adapted to their environment tend to survive and have better fitness. Natural selection is believed to be the motor of evolution. Evolution has occurred within all groups of organisms, including plant and animal populations.
Plants have evolved resistance in many circumstances, including resistance to pests and pathogens attacks. In response to insect herbivory attacks, plants have been using chemical defenses to resist. Consequently, to maximize their fitness, plant-feeding insects have co-evolved with plants to overcome plant defenses utilizing an array of strategies. Insects have evolved the ability to detoxify plant chemicals used for defenses and use the compounds as cues that favor the detection of the plant host. Insects have also evolved an adjusted sensory system allowing host cues detection and a nervous system that is able to integrate inputs from sensory neurons. The enhancement in the sensory and nervous systems allows the detection and avoidance of toxic plants as well as the excretion, sequestration, and degradation of plant toxins. Additionally, herbivory insects utilize target-site mutation, cuticular, humoral, and cellular defenses against plant chemical defenses. Moreover, insects have evolved to resist predation, parasitism, and pathogen attacks by means of a series of mechanisms, including cuticular adjustment, adaptive behavior, and chemical defenses.
With the intensive use of pesticides to manage agricultural pests, insect pests have evolved resistance to an array of insecticides using a variety of mechanisms. This type of evolution has been described as field-evolved resistance, which is a “genetically based decrease in susceptibility of a population to a toxin caused by exposure to the toxin in the field”. This is due to strong selection pressure that favors rapid evolution of resistance. For example, the widespread adoption of Bt crops in the U.S. has led to field-evolved resistance of corn earworm, also known as cotton bollworm, against Bt toxins. In some regions, Texas, for example, cotton bollworms being exposed to the Bt toxins in both corn and cotton throughout the year have been subjected to a high selection pressure, causing the pest to become quickly resistant to Bt toxins.
The use of beneficial arthropods to manage insects can favor a decrease in insecticide use and consequently reduce the selection pressure caused by pesticides. Although the evolution of resistance to predators and parasitoids tends to be prohibited by some factors (special and temporal refuges from enemies’ attacks, reciprocal evolution by control agents, and contrasting selection pressure from enemy species), the evolution of resistance to biological control agents has been reported for several insect pests including Argentine stem weevil, greater wax moth, and fruit fly. This is likely due to reduced plant and natural enemy diversity caused by intensive large-scale agriculture.
Several factors may play a role in the development of resistance. Large-scale homogenous agricultural systems do not allow enough refuges to sustain the susceptible strains, which would then mate with the resistant strains to dilute the resistance genes and maintain the susceptibility of the pest populations. Additionally, low biodiversity within the natural enemy population may favor the selection pressure. Coevolutionary arms races may play a significant role in that this may favor one participant in mutation and recombination rates.
Mechanisms of resistance:
Physiological resistance: Insects use physiological processes to become resistant to enemies and insecticides. In a pesticide use context, physiological resistance is defined as the capacity of an insect population to survive after being exposed to a concentration of insecticide that is known to be able to kill the totality of the population completely. However, the physiological process can also favor resistance against non-pesticide control methods. For instance, the fruit fly (Drosophila melanogaster) uses encapsulation to protect itself from koinobiont endoparasitoids. The encapsulation is a cellular immune response that follows three major stages, including the recognition of the parasitoid eggs as foreign, increasing the amount of circulating hemocytes that are produced by the lymph glands, and the lysis of the crystal cells allowing the release of prophenoloxidase which results in the melanization of the capsule surface.
Another way insects become resistant is through mutation in the target site of the toxicant. This physiological process can lead to resistance in insects against both plant defenses (toxic compounds released by the plant to protect itself from herbivory) and insecticides. This mutation can lead to target site insensitivity, meaning that even though the insect is being exposed to the toxic molecule, there will be no or reduced binding of the molecule to the target site, making the molecule ineffective. This mechanism of resistance is very common in many insecticide-resistant insect pests. Insects can also become resistant to toxic compounds from plants and to insecticides by evolving the ability to undergo detoxification of certain toxicants after exposure. This ability is also conferred by a series of mutations allowing the resistant insect to increase their enzyme production, which consequently increases their enzymatic activity and causes a rapid degradation of the toxicant into a nontoxic compound. This mechanism is also known as metabolic resistance.
Behavioral resistance: Many insect species have become resistant to certain host plants that use defense compounds to prevent herbivory through their plant selection and feeding behavior. For this behavior to occur, they must evolve the ability to detect toxic plants, which can be determined genetically or through a series of learning processes. Some other insects evolved the ability to deactivate or suppress the toxin produced by the plant hosts. For instance, the cotton bollworm uses its saliva, which is a gluco-oxidase, to cause a reduction in the level of nicotine produced in tobacco leaves. Other insects, when they feed on toxic plants, can excrete a significantly large amount of the accumulated toxic compound. Some insects even sequester the toxic compounds and use them for their own defense against predators and pathogens. Some insects that are hosts for parasitoids use a very effective behavioral resistant strategy by avoiding parasite contact or detection by choosing to niche away from the parasitoids or by choosing to locate themselves near a deterrent. Using this behavior, these insects are not directly resistant to the attackers but use what is present in their environment as tools to resist parasitism. Some other insects use alternative strategies, such as cryptic coloration or masquerade, to prevent their detection by predators and/or parasitoids. In this situation, they disguise themselves as something dangerous or unwanted to avoid being prayed on or parasitized.
Cuticular resistance: Insects depend heavily upon cuticular defenses to resist pathogens, parasitism, predation, and insecticides. For instance, to resist insecticide penetration, they develop a barrier in the outer layer of the cuticle either by changing the composition of the cuticle or by thickening it. This causes the toxicant to be penetrated slowly, consequently slowing the absorption of the contaminants to the insect bodies, where actions will take place.
Although the development of resistance is mostly beneficial for insects, there are some fitness costs associated with that. Physiological resistance, behavioral resistance, and cuticular resistance require the use of a large amount of energy; some energy that would have been allocated for growth, development, and reproduction is likely to be reduced, which would consequently reduce the fitness of the insect. Thus, fitness cost may cause an evolutionary constraint, which may reduce the rate or even prevent the evolution of resistance from occurring. Given that, an increase in resource availability is likely to favor the rate at which evolution occurs within a population.
In conclusion, resistance in insects can occur in a diversity of forms, and several factors may cause resistance to occur within insect populations. Additionally, while insect populations are more likely to be resistant to insecticide in large-scale agricultural systems, they can also become resistant to biological control agents, which underscores the importance of integrated pest management programs. The rate at which resistance occurs in a population closely depends on the intensity of selection pressure to which the insect populations are exposed. Thus, the more intense the selection pressure the quicker the populations will evolve resistant.
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