Ormia Ochracea
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''Ormia ochracea'' is a small yellow
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
in the family
Tachinidae The Tachinidae are a large and variable family of true flies within the insect order Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in this family ...
. It is notable for its parasitism of
crickets Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 8 ...
and its exceptionally acute directional hearing. The female is attracted to the song of the male cricket and deposits larvae on or around him, as was discovered in 1975 by the zoologist William H. Cade. ''Ormia ochracea'' is a
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
in
sound localization Sound localization is a listener's ability to identify the location or origin of a detected sound in direction and distance. The sound localization mechanisms of the mammalian auditory system have been extensively studied. The auditory system u ...
experiments because of its unique "ears", which are complex structures inside the fly's
prothorax The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
near the bases of its front legs. The fly is too small for the time difference of sound arriving at the two ears to be calculated in the usual way, yet it can determine the direction of sound sources with exquisite precision. The
tympanic membrane In the anatomy of humans and various other tetrapods, the eardrum, also called the tympanic membrane or myringa, is a thin, cone-shaped membrane that separates the external ear from the middle ear. Its function is to transmit sound from the air ...
s of opposite ears are directly connected mechanically, allowing resolution of nanosecond time differences and requiring a new
neural coding Neural coding (or Neural representation) is a neuroscience field concerned with characterising the hypothetical relationship between the stimulus and the individual or ensemble neuronal responses and the relationship among the electrical activit ...
strategy. Various research groups have designed low-noise differential microphones inspired by ''O. ochracea''’s directionally sensitive hearing system.


Distribution

''Ormia ochracea'' is native to the southeastern United States, including states such as
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
and
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. ''O. ochracea'' is also found throughout North America, South America, and the Caribbean, though its exact range is not known.


Life history

''Ormia ochracea'' has the full life cycle of egg, larvae, pupa, and adult. Once a female fly finds a suitable host, she deposits planidia (first
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
larvae) which then quickly burrow into the host. The
planidia A planidium is a specialized form of insect larva seen in the first-instar of a few families of insects that have parasitoidal ways of life. They are usually flattened, highly Sclerotization, sclerotized (hardened), and quite mobile. The functi ...
develop within the body of the field cricket host, embedding initially in muscle before migrating into the abdomen. The larvae molt within the host's abdomen and feed primarily on the host's muscle and fat. ''O. ochracea'' larvae typically complete development and emerge after about 7 days, which subsequently kills the host. The larvae pupate and emerge as adult flies approximately 2 weeks after emerging from the host.


Food resources

''O. ochracea'' is a
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
known to prey on several species of ''
Gryllus ''Gryllus'' is a genus of field cricket (Orthoptera, Gryllidae, Gryllinae). Members of the genus are typically 15–31 mm long and darkly coloured. The type species is ''Gryllus campestris'' L.: the European field cricket. Until the mid- ...
'' field crickets including ''
Gryllus integer ''Gryllus integer'', commonly known as the western trilling cricket, is one of many species of field cricket (subfamily Gryllinae) in the genus ''Gryllus''. It is called the "triller" field cricket because its song is nearly continuous rather tha ...
, Gryllus rubens, Gryllus texensis,'' and ''Gryllus firmus.'' Flies have been observed responding to various cricket songs, but seem to be limited to the family
Gryllidae The family ''Gryllidae'' contains the subfamilies and genera which entomologists now term true crickets. Having long, whip-like antennae, they belong to the Orthopteran suborder Ensifera, which has been greatly reduced in the last 100 years (''e ...
. The natural host of the fly may vary by location. Larvae of ''O. ochracea'' exhibit highest survival in its natural host and limited survival in other potential host species.


Host-finding

In 1975, William H. Cade experimentally demonstrated that ''Ormia ochracea'' uses the
mating call A mating call is the auditory signal used by animals to attract mates. It can occur in males or females, but literature is abundantly favored toward researching mating calls in females. In addition, mating calls are often the subject of mate choic ...
of the field cricket as a means to locate its host. Cade placed dead crickets on top of speakers playing cricket songs and various control sounds and recorded the amount of time the fly spent on either the control or test speaker. He found that flies spent more time on the speakers playing cricket songs and observed that the flies would always deposit larvae on and around the speaker which was playing cricket songs.


Learning

''O. ochracea'' have been shown to adjust their preference for host songs after exposure to different songs in the laboratory. In a 2011 study, flies that were previously exposed to the ''G. lineaticeps'' song chose the ''G. lineaticeps'' song over the ''G. integer'' song, and vice versa. This preference was very short term. ''O. ochraceas flexible learning capabilities may have been critical in expanding its host and geographical range.


Effects of infestation on host behavior

''O. ochracea'' infestation has been shown to affect the behavior and reproduction of host field crickets. Early in the infestation period, non-reproductive behavior is largely unimpaired because the parasitic larvae do not consume the digestive system or central nervous system of the host. After the larvae migrate to the host's abdomen, the host's mating, egg-laying, and fighting ability decline, most likely due to tissue damage caused by the larvae. Additionally, infestation of female crickets alters their mating preferences. ''
Gryllus lineaticeps ''Gryllus lineaticeps'', the variable field cricket, is a species of cricket in the subfamily Gryllinae. It is found in North America North America is a continent in the Northern Hemisphere and almost entirely within the Western ...
'' females normally prefer to respond to male songs with intermediate chirp rates over those with slow chirp rates, but females parasitized by ''O. ochracea'' show no preference between chirp rates. Reduced selectivity in infested female ''G. lineaticeps'' may be adaptive, as a female may be more likely to reproduce before being killed by the parasitoids if they are less selective.


Host defenses

Some species of
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by striki ...
which are parasitzed by ''O. ochracea'' have evolved methods to avoid infestation. For example, some members of the prey cricket ''
Teleogryllus oceanicus ''Teleogryllus oceanicus'', commonly known as the Australian, Pacific or oceanic field cricket, is a cricket found across Oceania and in coastal Australia from Carnarvon in Western Australia and Rockhampton in north-east Queensland Otte, D. & Ale ...
'' have a mutation called flat wing, in which the sound-producing structures of the male forewings are erased. The flat wing was first observed in 2003 on the
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
an island of
Kauai Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the 21st largest island ...
, and was also found on neighbouring
Oahu Oahu () (Hawaiian language, Hawaiian: ''Oʻahu'' ()), also known as "The Gathering place#Island of Oʻahu as The Gathering Place, Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over t ...
two years later. Genetic studies of crickets from each island show that the mutations arose from different genomic variations.


Enemies

Because field crickets commonly sing at night, ''O. ochracea'' are susceptible to predation by
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most ...
s. Studies have shown that ''O. ochracea'' has evolved an acoustic startle response to bat-like ultrasound, a response very similar to that of female crickets. ''O. ochracea'' has also been shown to demonstrate a sharp response boundary between the frequencies of cricket song and bat ultrasound.


Physiology


Directional hearing

In order for an animal to localize sound, it must be able to detect minute differences in intensity and time between the arrival of the sound to the ear closer to the source and the ear further from the source. ''O. ochracea'' displays a remarkable ability to localize sound despite the incredibly small distance (450-520μm) between its acoustic sensory organs. Its sound localization ability is facilitated by a cuticular structure which joins its ears, mechanically coupling their motion and magnifying interaural differences by a factor of about 20. Prior to ''O. ochracea'' no similar mechanism of auditory localization had been described.


Scientific significance

Several researchers have reported the construction of microphones inspired by the hearing system of ''O. ochracea.'' In 2009, R.N. Miles et al. designed and created a low-noise differential microphone inspired by the unique hearing system of ''O. ochracea'', for use in hearing aids''.'' The design of their microphone diaphragm, which measures 1 x 2 mm2, is based on the mechanically coupled ears of ''O. ochracea.'' Their microphone was found to have lower noise than commercially available hearing aid microphones while minimizing distance between sensors. In April 2015, a group from the
University of Strathclyde The University of Strathclyde ( gd, Oilthigh Shrath Chluaidh) is a public research university located in Glasgow, Scotland. Founded in 1796 as the Andersonian Institute, it is Glasgow's second-oldest university, having received its royal chart ...
and the MRC/CSO Institute for Hearing Research (IHR) announced that it had created a microphone based on ''O. ochraceas hearing system, and had been awarded a £430,000 grant by the U.K.
Engineering and Physical Sciences Research Council The Engineering and Physical Sciences Research Council (EPSRC) is a British Research Council that provides government funding for grants to undertake research and postgraduate degrees in engineering and the physical sciences, mainly to universi ...
to build and test the hearing aid for three years.


References


External links

*
NPR show on the hearing of ''Ormia ochracea''
* {{Taxonbar, from=Q3356515 Tachininae Taxa named by Jacques-Marie-Frangile Bigot Insects described in 1889 Diptera of North America