Strongyloides stercoralis

Ge­o­graphic Range

Strongy­loides ster­co­ralis is most no­tably found in the trop­ics and the sub­trop­ics, but it can occur in tem­per­ate cli­mates as well. Be­cause of the va­ri­ety of hosts which it can par­a­sitize, the worm can be found in var­i­ous areas around the world. In North Amer­ica, it has pri­mar­ily been found in large cities, specif­i­cally, New York City, Chicago, and Mon­treal. There is also a high in­ci­dence of Strongy­loides ster­co­ralis in Asia, Africa, trop­i­cal Amer­ica, the Pa­cific Is­lands, and even in parts of the for­mer So­viet Union. (Cheng, 1986; Roberts and Janovy, 2000)

Habi­tat

Free-liv­ing Strongy­loides ster­co­ralis pre­fer the moist soil of warm cli­mates. In the par­a­sitic gen­er­a­tion, the most com­mon habi­tat in the host is the mu­cosal lin­ing of the small in­tes­tine. Rarely, fi­lar­i­form lar­vae have been ob­served to take up res­i­dence in the lin­ing of the bronchi and tra­chea and lay their eggs there after trans­form­ing to adults. (Cheng, 1986)

  • Aquatic Biomes
  • lakes and ponds
  • rivers and streams
  • temporary pools

Phys­i­cal De­scrip­tion

Strongy­loides ster­co­ralis is gen­er­ally very long and cylin­dri­cal. It has four lay­ers of cu­ti­cle that are typ­i­cal of most ne­ma­todes. The epi­cu­ti­cle is the outer-most layer and the ex­o­cu­ti­cle is im­me­di­ately be­neath it. The meso­cu­ti­cle is be­neath the ex­o­cu­ti­cle and is gen­er­ally com­posed of very well or­ga­nized fi­brous lay­ers. The en­do­cu­ti­cle is the in­ner­most cu­ti­cle and al­though its makeup is sim­i­lar to the meso, its fibers are not well or­dered.

On the body sur­face, there are am­phids which act as chemore­cep­tors and they open to the out­side via pores which are lo­cated on the lips or on other an­te­rior ex­trem­i­ties. The worm has lat­eral alae, which can be seen as ridges which ex­tend, in some cases, across the body.

The mouth of Strongy­loides ster­co­ralis opens into a buc­cal cap­sule, which, in both sexes, is very small. From the buc­cal cap­sule, food moves into the esoph­a­gus which is very long, cylin­dri­cal, and lack­ing a bulb at its pos­te­rior end un­like most other ne­ma­todes. The esophagous is con­nected to the in­tes­tine via an esophago-in­testi­nal valve. The in­tes­tine is di­vided into three parts and is lined with a sin­gle layer of ep­ithe­lium. The most an­te­rior re­gion of the in­tes­tine is the ven­tric­u­lar re­gion and it has a se­cre­tory func­tion. The midre­gion, and the pos­te­rior pre­rec­tal re­gion func­tion pri­mar­ily in ab­sorp­tion. The in­tes­tine ends in the rec­tum and the al­i­men­tary tract even­tu­ally opens pos­te­ri­orly through the anus.

The Strongy­loides ster­co­ralis body cav­ity is known as a pseudo­coel. The re­pro­duc­tive sys­tem of males con­sists of vas def­er­ens which ex­tend from the testes and they en­large in the pos­te­rior por­tion of the body to form the sem­i­nal vesi­cle. Males pos­sess two cop­u­la­tory spicules in ad­di­tion to a gu­ber­nac­u­lum. Fe­males, on the other hand, usu­ally have two ovaries and an oviduct which ex­tends from it. At the ter­mi­nal end of the oviduct is a sem­i­nal re­cep­ta­cle, which con­nects the oviduct to the uterus. Free-liv­ing fe­males have uteri which con­tain more eggs than par­a­sitic fe­males. In par­a­sitic fe­males, the uteri are con­ver­gent and the vulva is in the pos­te­rior por­tion of the body. The free-liv­ing fe­male has a vulva that is more equa­to­r­ial.

Par­a­sitic adult fe­males range in length from 2.0 to 2.5 mm, lar­vae range in length from 300-800 mi­crom­e­ters, while the free-liv­ing male can grow up to 0.9 mm in length and 40 to 50 mi­crom­e­ters in width. (Cheng, 1986; Roberts and Janovy, 2000)

  • Sexual Dimorphism
  • female larger
  • sexes shaped differently
  • Range length
    0.9 to 2.5 mm
    0.04 to 0.10 in

De­vel­op­ment

There are three stages in the life-cy­cle of Strongy­loides ster­co­ralis: free-liv­ing, par­a­sitic, and au­toin­fec­tion. After cop­u­la­tion, the free-liv­ing fe­male ac­cu­mu­lates eggs in her uterus that con­tain par­tially de­vel­oped lar­vae. The eggs are laid in the soil where fur­ther de­vel­op­ment oc­curs. The rhabiti­form lar­vae ei­ther be­come free-liv­ing adults or they be­come in­fec­tive (fi­lar­i­form) 3rd stage lar­vae. Only fe­males can enter the next phase of de­vel­op­ment, the parastitic stage.

To con­tinue de­vel­op­ment, the fi­lar­i­form lar­vae must pen­e­trate a host's skin or be in­gested. If they enter the host through the skin, they must travel through host tis­sues in order to reach the final des­ti­na­tion, the in­tes­tine. There, the fe­males lay partheno­genetic eggs which give rise to rhabiti­form lar­vae. These even­tu­ally leave the host via the feces and de­velop into free-liv­ing adults or meta­mor­phose into fi­lar­i­form lar­vae.

An­other vari­ant of the life cycle is the au­toin­fec­tive phase. Fi­lar­i­form lar­vae, mi­grat­ing to the ex­te­rior, can re­in­fect the host by pen­e­trat­ing the skin and trav­el­ing in the blood to the lungs. (Cheng, 1986; Roberts and Janovy, 2000)

Re­pro­duc­tion

Fe­males may pro­duce a pher­momone to at­tract males. The male coils around a fe­male with his curved area over the fe­male gen­i­tal pore. The gu­ber­nac­u­lum, made of cu­ti­cle tis­sue, guides spicules which ex­tend through the cloaca and anus. Males use spicules to hold the fe­male dur­ing cop­u­la­tion. Ne­ma­tode sperm are amoe­boid-like and lack fla­gella.

There are three stages in the life-cy­cle of Strongy­loides ster­co­ralis: free-liv­ing, par­a­sitic, and au­toin­fec­tion. After cop­u­la­tion, the free-liv­ing fe­male ac­cu­mu­lates eggs in her uterus that con­tain par­tially de­vel­oped lar­vae. The eggs are laid in the soil where fur­ther de­vel­op­ment oc­curs. The rhabiti­form lar­vae ei­ther be­come free-liv­ing adults or they be­come in­fec­tive (fi­lar­i­form) 3rd stage lar­vae. Only fe­males can enter the next phase of de­vel­op­ment, the parastitic stage. (Barnes, 1987; Cheng, 1986; Roberts and Janovy, 2000)

  • Parental Investment
  • pre-fertilization
    • provisioning

Be­hav­ior

Rhabiti­form lar­vae of Strongy­loides ster­co­ralis molt four times be­com­ing sex­u­ally ma­ture adults. How­ever, if the ex­ter­nal en­vi­ron­ment be­comes un­fa­vor­able, they will meta­mor­phose into non-feed­ing fi­lar­i­form lar­vae that can in­fect hu­mans or some other host. As men­tioned ear­lier, if the fi­lar­i­form lar­vae enter the host through the skin, they must mi­grate to the in­tes­tine via the host's tis­sues. For years, it has been thought that the lar­vae are car­ried to the lungs by the blood. It is be­lieved that they in­habit the alve­oli by trav­el­ing through the right atrium and ven­tri­cle of the heart. The pul­monary artery even­tu­ally brings them to the lungs where they move up the bronchi and tra­chea and even­tu­ally are coughed up and swal­lowed. The fi­lar­i­form lar­vae then ma­ture in the in­tes­tine. This has only proven to be the case in hu­mans. In dogs the fi­lar­i­form lar­vae do not seem to travel through the lungs but seem to get to the in­tes­tine through any route.

Dur­ing the au­toin­fec­tive phase, fi­lar­i­form lar­vae reen­ter the host through the lower gut mu­cosa or the pe­ri­anal skin. (Cheng, 1986; Roberts and Janovy, 2000)

Com­mu­ni­ca­tion and Per­cep­tion

Ne­ma­todes within the Se­cer­nen­tea have phas­mids, which are uni­cel­lu­lar glands. Phas­mids likely func­tion as chemore­cep­tors. Fe­males may pro­duce pheromones to at­tract males.

Ne­ma­todes in gen­eral have papil­lae, setae and am­phids as the main sense or­gans. Setae de­tect mo­tion (mechanore­cep­tors), while am­phids de­tect chem­i­cals (chemore­cep­tors). (Barnes, 1987; Roberts and Janovy, 2000)

Food Habits

The three most im­por­tant de­fin­i­tive hosts for Strongy­loides ster­co­ralis are hu­mans, dogs, and cats. They fre­quently use other mam­mals as hosts as well, though with much less fre­quency. Par­a­sitic fe­males feed on the tis­sue of the host's in­ter­nal or­gans which in­cludes the in­testines as well as the lungs. Free-liv­ing adults and rhabiti­form lar­vae feed on or­ganic de­bris in soil or water.

Pha­ryn­geal glands and in­testi­nal ep­ithe­lium pro­duce di­ges­tive en­zymes to feed on the hosts’ body flu­ids. Ex­tra­cel­lu­lar di­ges­tion be­gins within the lumen and is fin­ished in­tra­cel­lu­larly. (Barnes, 1987; Cheng, 1986)

Pre­da­tion

These par­a­sites are prob­a­bly not preyed on di­rectly, but are in­gested. Lar­val mor­tal­ity is high as most of the par­a­sites do not reach ap­pro­pri­ate hosts.

Ecosys­tem Roles

The three most im­por­tant de­fin­i­tive hosts for Strongy­loides ster­co­ralis are hu­mans, dogs, and cats. They fre­quently use other mam­mals as hosts as well, though with much less fre­quency.

Species Used as Host

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Strongy­loides ster­co­ralis has no pos­i­tive ef­fects on hu­mans.

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Strongy­loides ster­co­ralis causes strongy­loidi­a­sis in hu­mans. In­fec­tion oc­curs via con­tact with fi­lar­i­form lar­vae in water or soil. The dis­ease is most preva­lent in areas of poor san­i­ta­tion such as those that have poor sewage dis­posal sites.

Strongy­loidi­a­sis oc­curs in three phases: in­va­sive, pul­monary, and in­testi­nal. Fi­lar­i­form lar­vae that pen­e­trate the skin cause itch­ing at the the entry site. Slight he­m­or­rhage and swelling may ap­pear as well and if bac­te­ria get into the site, in­flamm­tion is likely.

The pul­monary phase of the dis­ease is char­ac­ter­ized by wheez­ing, a burn­ing sen­sa­tion in the chest, spu­tum pro­duc­tion, as well as other symp­toms of bronchial pneu­mo­nia. Peo­ple who are thought to have a pul­monary in­fec­tion are di­ag­nosed with a chest x-ray. Pul­monary in­fec­tion can be fatal be­cause it can even­tu­ally re­sult in repi­ra­tory fail­ure. El­derly peo­ple, those with AIDS, and peo­ple who have had organ trans­plants have a very high risk of pul­monary strongy­loidi­a­sis. AIDS pa­tients and those with organ trans­plants are at such a high risk be­cause they are im­muno­sup­pressed, an ef­fect most likely due to cor­ti­cos­teroid use in the lat­ter group. The im­mune sys­tem is not able to fight off the par­a­site and this re­sults in a high rate of au­toin­fec­tion. Much care should be taken to di­ag­nose pul­monary strongy­loidi­a­sis prop­erly be­cause the symp­toms of it are very sim­i­lar to those of asthma and asthma is treated with cor­ti­cos­teroids, which could re­sult in in­creased au­toin­fec­tion.

The in­testi­nal phase oc­curs when the par­a­site pen­e­trates the in­testi­nal mu­cosa. Large patches of mu­cosa can be sloughed off in the process. An aching pain is felt in the ab­domen and in chronic cases, re­laps­ing col­i­tis is very likely. Ul­cer­a­tion of the in­tes­tine can occur in some cases if not treated prop­erly and this can lead to sep­ticemia which is, more often than not, fatal.

Strongy­loidi­a­sis is most often di­ag­nosed by a fecal smear, but in some cases, where in­fec­tion is not ram­pant, agar plate cul­ture to de­tect Strongy­loides ster­co­ralis is most ef­fec­tive (more so than the con­ven­tional fil­ter paper cul­ture). Some­times, em­bry­onated eggs may even be seen in the stool of pa­tients. How­ever, dif­fer­ent num­bers of ju­ve­niles pass through the feces each day and as au­toin­fec­tion oc­curs, the num­bers of ju­ve­niles ex­it­ing the body de­creases markedly.

There are three drugs that are pri­mar­ily used to treat strongy­loidi­a­sis. Each reg­i­men needs to be re­peated after one or two weeks be­cause it is dif­fi­cult to con­firm a pa­tient as cured. In a study con­ducted by Japan­ese doc­tors, iver­mectin proved to be most ef­fec­tive in treat­ing the dis­ease with a 97% rate of erad­i­ca­tion. It was given in a 6 mg sin­gle dose. The next most suc­cess­ful drug was al­ben­da­zole (400 mg/day for three days) with a cure rate of 77.4%. Pyrvinium pamoate once only found to cure about 23.3% of those tested. This drug was given at 5 mg/kg/day for 3 days. Thi­aben­da­zole was once thought to be an ef­fec­tive drug, but side ef­fects like nau­sea, vom­it­ing, and smelly urine have caused the drug to lose favor among doc­tors. (Roberts and Janovy, 2000; Ting, May 2000; Toma, et al., March 2000; Zaha, et al., Sept. 2000)

Con­ser­va­tion Sta­tus

There is no con­ser­va­tion sta­tus for Strongy­loides ster­co­ralis.

  • IUCN Red List [Link]
    Not Evaluated

Other Com­ments

It has pre­vi­ously been re­ported that Strongy­loides ster­co­ralis is amaz­ingly di­ver­gent from Strongy­loides ratti. ss­r­RNA se­quenc­ing found that there was only a 70% sim­i­lar­ity be­tween gene se­quences. This is un­usual be­cause two mem­bers of the same genus are never found to be that dif­fer­ent. The di­ver­gence has re­cently been found to be a re­sult of the fact that the gene se­quence of S. ster­co­ralis used in the study were hy­brids of ne­ma­tode and fun­gal se­quences. It has now been con­firmed that the ss­r­RNA be­tween the two species' is just over 90% sim­i­lar. (Dor­ris and Blax­ter, July 2000)

Con­trib­u­tors

Renee Sher­man Mul­crone (ed­i­tor).

Jason Hall­man (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor, Solomon David (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

agricultural

living in landscapes dominated by human agriculture.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

causes disease in humans

an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).

causes or carries domestic animal disease

either directly causes, or indirectly transmits, a disease to a domestic animal

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

cosmopolitan

having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

detritivore

an animal that mainly eats decomposed plants and/or animals

detritus

particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

freshwater

mainly lives in water that is not salty.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

marsh

marshes are wetland areas often dominated by grasses and reeds.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

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oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

parasite

an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

scrub forest

scrub forests develop in areas that experience dry seasons.

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

suburban

living in residential areas on the outskirts of large cities or towns.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

Ref­er­ences

Barnes, R. 1987. In­ver­te­brate Zo­ol­ogy. Or­lando, Florida: Dry­den Press.

Cheng, T. 1986. Gen­eral Par­a­sitol­ogy. Or­lando: Aca­d­e­mic Press.

Dor­ris, M., M. Blax­ter. July 2000. The small ri­bo­so­mal sub­unit RNA se­quence of Strongy­loides ster­co­ralis. In­ter­na­tional Jour­nal of Par­a­sitol­ogy, 30(8): 939-941.

Ohio State Uni­ver­sity, 2001?. "Strongy­loides ster­co­ralis" (On-line). Par­a­sites and Par­a­sito­log­i­cal Re­sources. Ac­cessed Sep­tem­ber 28, 2004 at Strongy­loides ster­co­ralis.

Roberts, L., J. Janovy. 2000. Foun­da­tions of Par­a­sitol­ogy. Boston: Mc­Graw Hill.

Ting, Y. May 2000. Pul­monary strongy­loidi­a­sis: Case re­port of two cases. Kaoh­si­ung J Med Sci, 16(5): 269-274.

Toma, H., Y. Sato, Y. Shi­roma, J. Kobayashi, I. Shimabukuro. March 2000. Com­par­a­tive stud­ies on the ef­fi­cacy of three an­thelminthics on treat­ment of human strongy­loidi­a­sis in Ok­i­nawa, Japan. SE Asian J Trop Med and Pub Health, 31(1): 147-151.

Zaha, O., T. Hi­rata, F. Kinjo, A. Saito. Sept. 2000. Strongy­loidi­a­sis: Progress in di­ag­no­sis and treat­ment. In­ter­nal Med­i­cine (Tokyo), 39(9): 695-700.