Entomophaga Grylli
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''Entomophaga grylli'' is a fungal
pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
which infects and kills
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are amongst what are possibly the most ancient living groups of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grassh ...
s. It is the causal agent of one of the most widespread diseases affecting grasshoppers. This is sometimes known as summit disease because infected insects climb to the upper part of a plant and grip the tip of the stem as they die; this ensures widespread dispersal of the fungal
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s. The fungus is a species complex with several different pathotypes, each one of which seems to be host-specific to different
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
of grasshoppers. The pathogen is being investigated for its possible use in
biological pest control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...
of grasshoppers.


Taxonomy

''Entomophaga grylli'' is a
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
which in North America includes the pathotypes ''E. macleodii'' and ''E. calopteni'', and in Australia ''E. praxibuli''. The different pathotypes can be distinguished by the formation or lack of formation of primary
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
, the size and the number of nuclei in the conidia, their
isozyme In biochemistry, isozymes (also known as isoenzymes or more generally as multiple forms of enzymes) are enzymes that differ in amino acid sequence but catalyze the same chemical reaction. Isozymes usually have different kinetic parameters (e.g. di ...
profiles, the host species involved, and the geographical range in which they are found. They all attack grasshoppers, but it is not known whether all grasshopper species are susceptible to infection. Each pathotype seems to be host-specific to certain grasshopper subfamilies.


Life cycle

''Entomophaga calopteni'' only produces
resting spore A resting spore is a resistant cell, adapted to survive adverse environmental conditions. Resting spore is a term commonly applied to both diatoms and fungi. In fungi and oomycetes A resting spore can be a spore created by fungi which is thickly ...
s, which are available to infect grasshoppers the following year. ''E. macleodii'' and ''E. praxibuli'' produce both resting spores and asexual conidia. Large numbers of conidia are produced under wet, humid conditions, and several cycles of infection can then occur in a single season. After landing on a potential host, a conidium produces a
germ tube A germ tube is an outgrowth produced by spores of spore-releasing fungi during germination. The germ tube differentiates, grows, and develops by mitosis to create somatic hyphae.C.J. Alexopolous, Charles W. Mims, M. Blackwell, ''Introductory My ...
which can grow through the cuticle into the
hemocoel In vertebrates, the circulatory system is a organ system, system of organs that includes the heart, blood vessels, and blood which is circulated throughout the body. It includes the cardiovascular system, or vascular system, that consists of ...
; once there, it produces amoeboid
protoplast Protoplast (), is a biology, biological term coined by Johannes von Hanstein, Hanstein in 1880 to refer to the entire cell, excluding the cell wall. Protoplasts can be generated by stripping the cell wall from plant, bacterium, bacterial, or f ...
s. These are cells without cell walls and seem to elude the insect's
immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...
. They multiply and kill the host. In some members of the species complex, they develop
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e with cell walls and grow through the cuticle, producing
conidiophore A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an Asexual reproduction, asexual, non-motility, motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word f ...
s and infective conidia. In these and other members of the species complex, they also have an intermediate stage producing resting spores with cell walls. After the insect
cadaver A cadaver, often known as a corpse, is a Death, dead human body. Cadavers are used by medical students, physicians and other scientists to study anatomy, identify disease sites, determine causes of death, and provide tissue (biology), tissue to ...
has fallen to the ground, the resting spores overwinter in the soil. A portion of them germinate in the spring, producing other spores that are forcibly ejected from the soil, landing on low vegetation where they come into contact with grasshoppers. They penetrate through the cuticle, proliferate and develop rapidly, the infected insect dying within about one week. At an advanced stage of the disease, an infected individual climbs to the top of a plant and dies with its limbs gripping the stem and its head pointing upwards. Some resting spores remain dormant in the soil for two or more years.
Epizootic In epizoology, an epizootic (or epizoötic, from Greek: ''epi-'' "upon" + ''zoon'' "animal") is a disease event in a nonhuman animal population analogous to an epidemic in humans. An epizootic disease (or ) may occur in a specific locale (an ...
outbreaks of disease in grasshopper populations in North America have been attributed to ''E. grylli'' but are usually localised and sporadic rather than widespread. They have been credited with ending many
outbreaks In epidemiology, an outbreak is a sudden increase in occurrences of a disease when cases are in excess of normal expectancy for the location or season. It may affect a small and localized group or impact upon thousands of people across an entire ...
of grasshopper species over the decades.


Use in biological control

In western Canada and the western United States, grasshoppers are estimated to cause over $400 million economic damage each year to crops and rangeland. From 1986 to 1992 an integrated pest management program was initiated by the
United States Department of Agriculture The United States Department of Agriculture (USDA) is an executive department of the United States federal government that aims to meet the needs of commercial farming and livestock food production, promotes agricultural trade and producti ...
and the
Animal and Plant Health Inspection Service The Animal and Plant Health Inspection Service (APHIS) is an agency of the United States Department of Agriculture (USDA) based in Riverdale Park, Maryland, Riverdale, Maryland responsible for protecting animal health, animal welfare, and plant h ...
to attempt to control grasshopper numbers without the use of vast quantities of insecticide. The inclusion of the ''E. grylli'' complex in the program was investigated. A disadvantage to its use is that, unlike entomopathogens such as '' Metarhizium acridum'', the fungus cannot be mass-produced and its effectiveness depends on the weather conditions (more grasshoppers are infected in warm, moist conditions). Attempts to control grasshoppers with this fungus have been largely ineffective; insects can be successfully infected by injecting them with the pathogen, but introduction of North American pathotypes into Australia and vice versa have failed to establish long term infections. The pathogen has potential for biological control of grasshoppers but more research is needed.


References

{{Taxonbar, from=Q6709410 Entomophthorales Fungus species Parasitic fungi