Entomopathogenic nematode

Entomopathogenic nematodes are a group of nematodes (thread worms), causing death to insects. The term entomopathogenic has a Greek origin entomon, refers to insect, and pathogenic, which denotes causing disease. They are multi-cellular metazoans that occupy a bio control middle ground between microbial pathogens and predator/ parasitoids, and are habitually grouped with pathogens, most likely because of their symbiotic relationship with bacteria. Although many other parasitic thread worms cause diseases in living organisms (sterilizing or otherwise debilitating their host), entomopathogenic nematodes, are specific in only infecting insects. Entomopathogenic nematodes (EPNs) live parasitically inside the infected insect host, and so they are termed as endoparasitic. They infect many different types of insects living in the soil like the larval forms of moths, butterflies, flies and beetles as well as adult forms of beetles, grasshoppers and crickets. EPNs have been found in all over the world and a range of ecologically diverse habitats. They are highly diverse, complex and specialized. The most commonly studied entomopathogenic nematodes are those that can be used in the biological control of harmful insects, the members of Steinernematidae and Heterorhabditidae (Gaugler 2006). They are the only insect-parasitic nematodes possessing an optimal balance of biological control attributes. (Cranshaw & Zimmerman 2013). Entomopathogenic nematodes are a group of nematodes (thread worms), causing death to insects. The term entomopathogenic has a Greek origin entomon, refers to insect, and pathogenic, which denotes causing disease. They are multi-cellular metazoans that occupy a bio control middle ground between microbial pathogens and predator/ parasitoids, and are habitually grouped with pathogens, most likely because of their symbiotic relationship with bacteria. Although many other parasitic thread worms cause diseases in living organisms (sterilizing or otherwise debilitating their host), entomopathogenic nematodes, are specific in only infecting insects. Entomopathogenic nematodes (EPNs) live parasitically inside the infected insect host, and so they are termed as endoparasitic. They infect many different types of insects living in the soil like the larval forms of moths, butterflies, flies and beetles as well as adult forms of beetles, grasshoppers and crickets. EPNs have been found in all over the world and a range of ecologically diverse habitats. They are highly diverse, complex and specialized. The most commonly studied entomopathogenic nematodes are those that can be used in the biological control of harmful insects, the members of Steinernematidae and Heterorhabditidae (Gaugler 2006). They are the only insect-parasitic nematodes possessing an optimal balance of biological control attributes. (Cranshaw & Zimmerman 2013). Because of their economic importance, the life cycles of the genera belonging to families Heterorhabditidae and Steinernematidae are well studied. Although not closely related, phylogenetically, both share similar life histories (Poinar 1993). The cycle begins with an infective juvenile, whose only function is to seek out and infect new hosts. When a host has been located, the nematodes penetrate into the insect body cavity, usually via natural body openings (mouth, anus, spiracles) or areas of thin cuticle. (Shapiro-Ilan, David I., and Randy Gaugler. 'Nematodes.') After entering an insect, infective juveniles release an associated mutualistic bacterium from their gut which multiplies rapidly. These bacteria of the genus Xenorhabdus or Photorhabdus, for steinerernematides and heterorhabditids, respectively—cause host mortality within 24-48 hours. The nematodes provide shelter to the bacteria, which, in return, kill the insect host and provide nutrients to the nematode. Together, the nematodes and bacteria feed on the liquefying host, and reproduce for several generations inside the cadaver maturing through the growth stages of J2-J4 into adults. Steinernematids infective juveniles may become males or females, whereas heterorhabditids develop into self-fertilizing hermaphrodites with later generations producing two sexes. When food resources in the host become scarce, the adults produce new infective juveniles adapted to withstand the outside environment. The life cycles of the EPNs are completed within a few days.(Shapiro-Ilan, David I., and Randy Gaugler. 'Nematodes.') After about a week, hundreds of thousands of infective juveniles emerge and leave in search of new hosts, carrying with them an inoculation of mutualistic bacteria, received from the internal host environment (Boemare 2002, Gaugler 2006). Their growth and reproduction depends upon conditions established in the host cadaver by the bacterium. The nematodes bacterium contributes anti-immune proteins to assist in overcoming their host defenses.(Shapiro-Ilan, David I., and Randy Gaugler. 'Nematodes.') The foraging strategies of entomopathogenic nematodes vary between species, influencing their soil depth distributions and host preferences. Infective juveniles use strategies to find hosts that vary from ambush and cruise foraging (Campbell 1997). In order to ambush prey, some Steinernema species nictate, or raise their bodies off the soil surface so they are better poised to attach to passing insects, which are much larger in size (Campbell and Gaugler 1993). Many Steinernema are able to jump by forming a loop with their bodies that creates stored energy which, when released, propels them through the air (Campbell and Kaya 2000). Other species adopt a cruising strategy and rarely nictate. Instead, they roam through the soil searching for potential hosts. These foraging strategies influence which hosts the nematodes infect. For example, ambush predators such as Steinernema carpocapsae infect more insects on the surface, while cruising predators like Heterorhabditis bacteriophora infect insects that live deep in the soil (Campbell and Gaugler 1993). Inside their insect hosts, EPNs experience both intra and interspecific competition. Intraspecific competition takes place among nematodes of the same species when the number of infective juveniles penetrating a host exceeds the amount of resources available. Interspecific competition occurs when different species compete for resources. In both cases, the individual nematodes compete with each other indirectly by consuming the same resource, which reduces their fitness and may result in the local extinction of one species inside the host (Koppenhofer and Kaya 1996). Interference competition, in which species compete directly, can also occur. For example, a steinernematid species that infects a host first usually excludes a heterorhabditid species. The mechanism for this superiority may be antibiotics produced by Xenorhabdus, the symbiotic bacterium of the steinernematid. These antibiotics prevent the symbiotic bacterium of the heterorhabditid from multiplying (Kaya and Koppenhofer1996). In order to avoid competition, some species of infective juveniles are able to judge the quality of a host before penetration. The infective juveniles of S. carpocapsae are repelled by 24-hour-old infections, likely by the smell of their own species’ mutualistic bacteria (Grewal et al. 1997). Interspecific competition between nematode species can also occur in the soil environment outside of hosts. Millar and Barbercheck (2001) showed that the introduced nematode Steinernema riobrave survived and persisted in the environment for up to a year after its release. S. riobrave significantly depressed detection of the endemic nematode H. bacteriophora, but never completely displaced it, even after two years of continued introductions. S. riobrave had no effect on populations of the native nematode, S. carpocapsae, though, which suggests that coexistence is possible. Niche differentiation appears to limit competition between nematodes. Different foraging strategies allow two species to co-exist in the same habitat. Different foraging strategies separate the nematodes in space and enable them to infect different hosts. EPNs also occur in patchy distributions, which may limit their interactions and further support coexistence (Kaya and Koppenhofer 1996). Entomopathogenic nematodes are typically found in patchy distributions, which vary in space and time, although the degree of patchiness varies between species (reviewed in Lewis 2002). Factors responsible for this aggregated distribution may include behavior, as well as the spatial and temporal variability of the nematodes natural enemies, like nematode trapping fungus. Nematodes also have limited dispersal ability. Many infective juveniles are produced from a single host which could also produce aggregates. Patchy EPN distributions may also reflect the uneven distribution of host and nutrients in the soil (Lewis et al. 1998; Stuart and Gaugler 1994; Campbell et al. 1997, 1998). EPNs may persist as metapopulations, in which local population fragments are highly vulnerable to extinction, and fluctuate asynchronously (Lewis et al. 1998). The metapopulation as a whole can persist as long as the rate of colonization is greater or equal to the rate of population extinction (Lewis et al. 1998). The founding of new populations and movement between patches may depend on the movement of infective juveniles or the movement of infected hosts (Lewis et al. 1998). Recent studies suggest that EPNs may also use non-host animals, such as isopods and earthworms for transport (Eng et al.2005, Shapiro et al. 1993) or can be scavengers (San-Blas and Gowen, 2008). Parasites can significantly affect their hosts, as well as the structure of the communities to which they and their hosts belong (Minchella and Scott 1991). Entomopathogenic nematodes have the potential to shape the populations of plants and host insects, as well as the species composition of the surrounding animal soil community. Entomopathogenic nematodes affect populations of their insect hosts by killing and consuming individuals. When more EPNs are added to a field environment, typically at concentrations of 250,000 individuals per square metre, the population of host insects measurably decreases (Campbell et al. 1998, Strong et al. 1996). Agriculture exploits this finding, and the inundative release of EPNs can effectively control populations of soil insect pests in citrus, cranberries, turfgrass, and tree fruit (Lewis et al. 1998).If entomopathogenic nematodes suppress the population of insect root herbivores, they indirectly benefit plants by freeing them from grazing pressure. This is an example of a trophic cascade in which consumers at the top of the food web (nematodes) exert an influence on the abundance of resources (plants) at the bottom. The idea that plants can benefit from the application of their herbivore’s enemies is the principle behind biological control. Consequently, much of EPN biological research is driven by agricultural applications. Examples of the top-down effects of entomopathogenic nematodes are not restricted to agricultural systems. Researchers at the Bodega Marine Laboratory examined the strong top-down effects that naturally occurring EPNs can have on their ecosystem (Strong et al. 1996). In a coastal shrubland food chain the native EPN, Heterorhabditis heplialus, parasitized ghost moth caterpillars, and ghost moth caterpillars consumed the roots of bush lupine. The presence H. heplialus correlated with lower caterpillar numbers and healthier plants. In addition, the researchers observed high mortality of bush lupine in the absence of EPNs. Old aerial photographs over the past 40 years indicated that the stands where nematodes were prevalent had little or no mass die-off of lupine. In stands with low nematode prevalence, however, the photos showed repeated lupine die-offs. These results implied that the nematode, as a natural enemy of the ghost moth caterpillar, protected the plant from damage. The authors even suggested that the interaction was strong enough to affect the population dynamics of bush lupine (Strong et al. 1996).